CHAPTER 4-6 INVERTEBRATES: ROTIFER TAXA

available at <www.bryoecol.mtu.edu>.

CHAPTER 4-6

INVERTEBRATES: ROTIFER TAXA

TABLE OF CONTENTS

Taxa

on

Bryophytes ... 4-6-2

CLASS

BDELLOIDEA ... 4-6-6

Adinetidae... 4-6-6

Habrotrochidae... 4-6-7

Philodinidae ... 4-6-8

CLASS

MONOGONONTA ... 4-6-10

Order

Collothecacea... 4-6-10

Collothecidae ... 4-6-10

Order

Flosculariacea ... 4-6-13

Conochilidae ... 4-6-13

Filiniidae ... 4-6-13

Flosculariidae... 4-6-14

Hexarthriidae ... 4-6-15

Testudinellidae... 4-6-16

Order

Ploimida... 4-6-17

Brachionidae ... 4-6-18

Dicranophoridae... 4-6-20

Epiphanidae ... 4-6-24

Euchlanidae... 4-6-25

Gastropodidae ... 4-6-27

Lecanidae... 4-6-27

Ituridae... 4-6-34

Summary ... 4-6-34

Acknowledgments... 4-6-34

Literature Cited ... 4-6-35

CHAPTER 4-6

INVERTEBRATES: ROTIFER TAXA

Figure 1. Rotifer on a Sphagnum leaf. Photo by Marek Miś at <http://www.mismicrophoto.com/>.

Taxa on Bryophytes

With about 2200 species, rotifers are a group with a

wide range of aquatic, marine, and limnoterrestrial species,

permitting us to analyze habitat relations. This is not true

with respect to bryophytes because few studies describe

those in the bryophyte habitat, and those that do typically

simply indicate "moss." This is demonstrated by the

delineation of rotifer habitats in the comprehensive study

on the relationship of rotifers to habitat, using only

macrophytes (housing periphytic rotifers), open water

(with planktonic forms), minerogenous sediments (with

psammon and hyporheos),

organogenous sediments, and

other organisms (

i.e.

parasites and epizoans) (Pejler 1995).

Bryophytes are not given separate attention. Pejler (1995)

pointed out that rotifers are mostly cosmopolitan, hence

suggesting that ecological barriers are more important in

determining their distribution. Nevertheless, Pejler

considers rotifers to lack strong restrictions of habitat.

Extreme environments do support few species, but large

numbers of individuals, typically primary consumers. On

the other hand, when rotifer species are numerous the

differences in their morphology are so great that patterns of

adaptations are difficult to define.

The few adaptations that do exist include protection

from predation among planktonic species. Differences in

structure of the trophus seem to facilitate differences in

food type. Even in extreme environments, the differences

don't seem to correlate with the habitat and the closest

relatives seem to occur in "normal" habitats. Pejler

considered that adaptations to chemical and physical

environments may develop rapidly in geologic time,

whereas those changes that are more fundamental occur

over a longer time period. It is this group where changes in

trophi are most apparent.

Although many taxa can be found on bryophytes

(Table 1), few have been studied relative to bryophytes,

and finding the existing studies among published literature

can be a bit hit or miss. I am unable to summarize

adaptations except to suggest that being small (which

applies to the entire phylum) and being able to attach may

be advantages. The trophi need to be adapted to the

available food, with detritus being abundant among the

bryophytes. The list provided here is not intended to be

comprehensive and the ecological information included

with the images is very incomplete. Likewise, the

distribution of species is poorly known, although many are

considered cosmopolitan. I have indicated cosmopolitan

where I found references so-stating, but I have rarely been

able to ascertain countries or distributions and thus have

not included those. Due to these limitations, these chapters

are organized by classification rather than ecology.

Table 1. Species and genera of rotifers known from collections of bryophytes or in bog pools. Authors indicate those who have reported the rotifer species in a collection of bryophytes or from a Sphagnum pool. Those indicated by * indicate those species that

have been collected on Sphagnum; + indicates that those collected from Sphagnum were also collected from other bryophytes. If no superscript is given, the author/collector simply said moss. An indication of bog refers to a Sphagnum bog, but not necessarily on a moss (and possibly not a true bog). Please note that some, perhaps most, of the rotifers in this list may not be true bryophyte dwellers, but rather occasional visitors. Those species that have been found in more than one location in association with bryophytes have the species name in bold as this may be an indication it is more than an occasional visitor. Nomenclature follows Segers 2007.

Adineta barbata* – S. subsecundum Horkan 1931;

Hingley 1993; Jersabek et al. 2003

Adineta gracilis* Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Adineta steineri Hirschfelder et al. 1993

Adineta vaga Hingley 1993 Adineta tuberculosa Horkan 1931

Albertia naidis* – Fontinalis Pejler & Bērziņš 1993;

Jersabek et al. 2003

Anuraeopsis fissa Horkan 1931

Aspelta angusta* – Fontinalis Pejler & Bērziņš 1993;

Jersabek et al. 2003

Aspelta aper*+ – Fontinalis Pejler & Bērziņš 1993 Aspelta beltista* Jersabek et al. 2003 Aspelta chorista* Jersabek et al. 2003

Aspelta circinator* Horkan 1931; Hingley 1993; – Fontinalis Pejler & Bērziņš 1993; Jersabek et al. 2003

Brachionus urceolaris Hingley 1993 Bradyscela clauda Madaliński 1961

Bryceella perpusilla – terrestrial mosses Wilts et al. 2010

Bryceella stylata

*

Bryceella tenella* Hingley 1993; Jersabek et al. 2003

Bryceella voigti Hingley 1993 Callidena symbiotica Hudson 1889

Cephalodella anebodica – bogs Błedzki & Ellison 2003

Cephalodella apocoela* Hingley 1993; Jersabek et al. 2003

Cephalodella auriculata Hingley 1993 Cephalodella belone Jersabek et al. 2003

Cephalodella biungulata

*

Cephalodella catellina Horkan 1931; Hingley 1993

Cephalodella compressa Jersabek et al. 2003 Cephalodella dorseyi –

Fontinalis

*

Cephalodella eva Horkan 1931 Cephalodella exigua Jersabek et al. 2003

Cephalodella forficula Horkan 1931; Hingley 1993 Cephalodella gibba* Horkan 1931; Hingley 1993; De Smet 2001; Jersabek et al. 2003 Cephalodella gracilis Madaliński 1961

Cephalodella hoodii Horkan 1931 Cephalodella inquilina Jersabek et al. 2003

Cephalodella intuta Hingley 1993 Cephalodella lepida – bog Jersabek et al. 2003

Cephalodella licinia

*

Cephalodella megalotrocha Horkan 1931 Cephalodella mira* Jersabek et al. 2003

Cephalodella mucronata

*

Cephalodella nana Hingley 1993 Cephalodella nelitis

*

Cephalodella pheloma Hingley 1993

Cephalodella physalis – bog Hingley 1993; Jersabek et al. 2003

Cephalodella rostrum Hingley 1993

Cephalodella sterea Horkan 1931 Cephalodella subsecundum Jersabek et al. 2003

Cephalodella tachyphora Jersabek et al. 2003

Cephalodella tantilla Hingley 1993 Cephalodella tantilloides Hingley 1993 Cephalodella ventripes Hingley 1993

Ceratotrocha cornigera Horkan 1931; Hingley 1993

Collotheca ambigua – sessile on Sphagnum Hingley 1993

Collotheca annulata – sessile on Sphagnum Hingley 1993

Collotheca calva – sessile on Sphagnum Hingley 1993

Collotheca campanulata – sessile on Sphagnum Hingley 1993

Collotheca catellina Jersabek et al. 2003 Collotheca coronetta – sessile on Sphagnum Hingley 1993

Collotheca crateriformis* Jersabek et al. 2003 Collotheca heptabrachiata Edmondson 1940

Collotheca hoodii – sessile on Sphagnum Hingley 1993

Collotheca ornata – sessile on Sphagnum Hingley 1993

Collotheca quadrinodosa – sessile on Sphagnum Hingley 1993

Collotheca spinata – sessile on Sphagnum Hingley 1993

Collotheca trilobata – sessile on Sphagnum Hingley 1993 Colurella adriatica Horkan 1931; Hingley 1993

Colurella colurus Madaliński 1961 Colurella hindenburgi* – S. subsecundum Jersabek et al. 2003

Colurella obtusa Horkan 1931; Hingley 1993

Colurella obtusa clausa – bogs Błedzki & Ellison 2003

Colurella paludosa Hingley 1993

Colurella tessellata – bogs Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Conochilus – sessile on Sphagnum Hingley 1993

Cyrtonia tuba Horkan 1931 Dicranophorus alcinus* Jersabek et al. 2003

Dicranophorus artamus* Jersabek et al. 2003 Dicranophorus biastis* Jersabek et al. 2003 Dicranophorus capucinus* Jersabek et al. 2003 Dicranophorus colastes* Jersabek et al. 2003 Dicranophorus corystis* Jersabek et al. 2003 Dicranophorus edestes – Fontinalis Jersabek et al. 2003 Dicranophorus epicharus* Pejler & Bērziņš 1993 Dicranophorus facinus* Jersabek et al. 2003

Dicranophorus forcipatus Horkan 1931;

Fontinalis Pejler & Bērziņš 1993

Dicranophorus haueri – Fontinalis Pejler & Bērziņš 1993

Dicranophorus hercules Hingley 1993 Dicranophorus isothes* Jersabek et al. 2003

Dicranophorus lenapensis – Fontinalis Jersabek et al. 2003

Dicranophorus longidactylum Hingley 1993

Dicranophorus luetkeni* Hingley 1993;

Pejler & Bērziņš 1993; Jersabek et al. 2003

Dicranophorus robustus Hingley 1993; Pejler & Bērziņš 1993

Dicranophorus robusta europaeus – Fontinalis

Pejler & Bērziņš 1993

Dicranophorus rostratus* Hingley 1993; Jersabek et al. 2003 Dicranophorus saevus* Jersabek et al. 2003

Dicranophorus spiculatus - Fontinalis Jersabek et al. 2003 Dicranophorus thysanus – Sphagnum bog & pond

Jersabek et al. 2003

Dicranophorus uncinatus Horkan 1931; Hingley 1993; Fontinalis Pejler & Bērziņš 1993

Didymodactylos Ricci & Melone 2000

Dipleuchanis paludosa Hingley 1993 Dipleuchanis propatula Hingley 1993

Dissotrocha aculeata* Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Dissotrocha macrostyla Horkan 1931; Hingley 1993

Dissotrocha spinosa Hingley 1993 Dorria dalecarlica – submerged moss on rock

Jersabek et al. 2003

Elosa worrallii

*

Encentrum arvicola* Pejler & Bērziņš 1993 Encentrum carlini* Jersabek et al. 2003 Encentrum elongatum* Pejler & Bērziņš 1993 Encetrum eurycephalum – Fontinalis Pejler & Bērziņš 1993

Encentrum felis* Hingley 1993; Jersabek et al. 2003 Encetrum fluviatile – Fontinalis Pejler & Bērziņš 1993 Encentrum glaucum Hingley 1993 Encentrum incisum* Pejler & Bērziņš 1993 Encetrum lupus* – Fontinalis Pejler & Bērziņš 1993

Encentrum mustela Hingley 1993;

Fontinalis Pejler & Bērziņš 1993

Encentrum sutor* Pejler & Bērziņš 1993 Encentrum sutoroides* Pejler & Bērziņš 1993 Encentrum tobyhannaensis* Jersabek et al. 2003 Encentrum tyrphos* Pejler & Bērziņš 1993

Enteroplea lacustris* Horkan 1931; Jersabek et al. 2003

Eosphora ehrenbergi Horkan 1931 Eosphora najas Madaliński 1961

Eothinia elongata Horkan 1931 Euchlanis callysta* Jersabek et al. 2003

Euchlanis calpidia* Jersabek et al. 2003 Euchlanis dilatata Jersabek et al. 2003

Euchlanis incisa Hingley 1993 Euchlanis meneta Hingley 1993 Euchlanis parva Hingley 1993 Euchlanis proxima Hingley 1993 Euchlanis pyriformis Horkan 1931

Euchlanis triquetra – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Euchlanis triquetra subsp pellucida Jersabek et al. 2003

Filinia longiseta Horkan 1931

Filinia terminalis – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Floscularia conifera – sessile on Sphagnum Hingley 1993 Gastropus hyptopus Horkan 1931; Hingley 1993 Gastropus minor – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Habrotrocha ampulla* Jersabek et al. 2003

Habrotrocha angusticollis

*

Habrotrocha aspera Horkan 1931 Habrotrocha bidens Hingley 1993

Habrotrocha collaris Horkan 1931; Hingley 1993 Habrotrocha constricta Horkan 1931; Hingley 1993

Habrotrocha elegans Hingley 1993 Habrotrocha eremita Peters et al. 1993 Habrotrocha flava Hirschfelder et al. 1993 Habrotrocha fusca Hirschfelder et al. 1993 Habrotrocha insignis Hirschfelder et al. 1993

Habrotrocha lata* Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Habrotrocha longula Hingley 1993 Habrotrocha microcephala Madaliński 1961

Habrotrocha milnei Hingley 1993 Habrotrocha minuta Hingley 1993

Habrotrocha pulchra Horkan 1931 Habrotrocha pusilla Horkan 1931

Habrotrocha reclusa* – S. subsecundum Hingley 1993;

Jersabek et al. 2003

Habrotrocha roeperi* Horkan 1931; Hingley 1993

Habrotrocha rosa Madaliński 1961 Habrotrocha tridens Madaliński 1961

Hexarthra mira Horkan 1931 Itura aurita Horkan 1931 Kellicottia longispina Madaliński 1961

Keratella mixta* Jersabek et al. 2003

Keratella quadrata Hingley 1993

Keratella serrulata* Bērziņš & Pejler 1987; Hingley 1993 Lecane agilis* Hingley 1993; Jersabek et al. 2003 Lecane calcaria* Jersabek et al. 2003

Lecane clara Hingley 1993 Lecane climacois Jersabek et al. 2003

Lecane closterocerca Hingley 1993 Lecane cornuta Hingley 1993 Lecane curvicornis acronyrha - Sphagnum bog

Jersabek et al. 2003

Lecane depressa – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Lecane elasma Jersabek et al. 2003

Lecane flexilis Hingley 1993;

Riccia fluitans Jersabek et al. 2003

Lecane galeata* – Sphagnum bog Hingley 1993;

S. subsecundum Jersabek et al. 2003 Lecane gallagherorum* Jersabek et al. 2003

Lecane hamata Hingley 1993

Lecane inermis* Hingley 1993; Jersabek et al. 2003 Lecane lauterborni Jersabek et al. 2003 Lecane ligona Jersabek et al. 2003

Lecane lunaris Madaliński 1961; Hingley 1993

Lecane mira* Jersabek et al. 2003 Lecane mitis* Jersabek et al. 2003 Lecane pertica Jersabek et al. 2003

Lecane pyrrha* – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Lecane rhopalura – submerged moss Jersabek et al. 2003

Lecane quadridentata Horkan 1931 Lecane satyrus* Jersabek et al. 2003

Lecane scutata Koste & Shiel 1990

Lecane signifera* Hingley 1993; Jersabek et al. 2003

Lecane signifera ploenensis*Hingley 1993; Jersabek et al. 2003

Lecane stichaea* Hingley 1993; Jersabek et al. 2003 Lecane subulata Jersabek et al. 2003 Lecane tenuiseta* Jersabek et al. 2003 Lecane thalera* Jersabek et al. 2003 Lecane tryphema – Sphagnum bog Jersabek et al. 2003 Lecane ungulata Madaliński 1961

Lepadella acuminata Hingley 1993 Lepadella akrobeles* Jersabek et al. 2003

Lepadella bractea* Jersabek et al. 2003 Lepadella eurysterna – Fontinalis novae-angliae

Jersabek et al. 2003

Lepadella ovalis Hingley 1993 Lepadella patella Hingley 1993 Lepadella pterygoidea* Jersabek et al. 2003

Lepadella pterygoides Hingley 1993

Lepadella triba* Hingley 1993; Jersabek et al. 2003

Lepadella triptera Horkan 1931; Hingley 1993

Lepadella venefica* – emersed S. subsecundum;

Sphagnum bog Jersabek et al. 2003 Lindia annecta de Manuel Barrabin 2000

Lindia torulosa Hingley 1993

Macrochaetus collinsi Hingley 1993 Macrochaetus multispinosus* Jersabek et al. 2003

Macrotrachela bilfingeri Madaliński 1961

Macrotrachela ehrenbergii* Peters et al. 1993;

Jersabek et al. 2003

Macrotrachela habita* Horkan 1931; Hirschfelder et al. 1993;

Jersabek et al. 2003

Macrotrachela insolita Hirschfelder et al. 1993

Macrotrachela multispinosa Horkan 1931; Hingley 1993 Sphagnum bog; on "tree moss" Jersabek et al. 2003

Macrotrachela muricata Horkan 1931 Macrotrachela musculosa Hirschfelder et al. 1993

Macrotrachela nana Madaliński 1961

Macrotrachela papillosa Horkan 1931; Hingley 1993 Macrotrachela plicata Horkan 1931; Hingley 1993;

on "tree moss" Jersabek et al. 2003 Macrotrachela punctata Hirschfelder et al. 1993

Macrotrachela quadricornifera* Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Macrotrachela zickendrahti* Jersabek et al. 2003

Mikrodides chalaena Horkan 1931

Microcodon clavus* Horkan 1931; Hingley 1993;

Jersabek et al. 2003

Mniobia incrassata Hingley 1993

Mniobia magna Hingley 1993 Mniobia obtuscicornis Hingley 1993 Mniobia orta Peters et al. 1993

Mniobia russeola Horkan 1931; Hirschfelder et al. 1993 Mniobia scarlatina – "tree moss" Jersabek et al. 2003

Mniobia symbiotica Horkan 1931; Hingley 1993

Mniobia tetraodon Horkan 1931

Monommata actices* Hingley 1993; Jersabek et al. 2003

Monommata aequalis Horkan 1931

Monommata aeschyna

*

Monommata longiseta Hingley 1993 Monommata maculata Hingley 1993 Monommata phoxa Hingley 1993 Mytilina macrocera* Jersabek et al. 2003

Mytilina mucronata Horkan 1931; Hingley 1993

Mytilina ventralis var. brevispina Hingley 1993

Notommata allantois Hingley 1993 Notommata brachyota Horkan 1931

Notommata cerberus Horkan 1931; Hingley 1993

Notommata cherada – Sphagnum bog Jersabek et al. 2003

Notommata contorta Hingley 1993

Sphagnum pool Jersabek et al. 2003

Notommata copeus Horkan 1931; Hingley 1993

Notommata cyrtopus Horkan 1931

Notommata falcinella* Hingley 1993;

Sphagnum subsecundum Jersabek et al. 2003 Notommata fasciola* Jersabek et al. 2003

Notommata groenlandica Hingley 1993

Notommata pachyura Horkan 1931; Hingley 1993

Notommata saccigera Hingley 1993

Notommata tripus Horkan 1931; Hingley 1993

Otostephanos macrantennus Ricci 1998 Otostephanos regalis Hirschfelder et al. 1993 Otostephanos torquatus Peters et al. 1993

Paracolurella aemula* Jersabek et al. 2003 Paracolurella logima* Jersabek et al. 2003 Pedipartia gracilis* Jersabek et al. 2003

Philodina acuticornis Hingley 1993 Philodina brevipes Hingley 1993 Philodina citrina Hirschfelder et al. 1993;

Sphagnum bog; "tree moss" Jersabek et al. 2003

Philodina erythrophthalma Horkan 1931

Philodina flaviceps Horkan 1931; Madaliński 1961

Philodina nemoralis Hingley 1993

Philodina plena* Hirschfelder et al. 1993;

Jersabek et al. 2003

Philodina roseola Hirschfelder et al. 1993

Philodina rugosa Horkan 1931; Hingley 1993

Philodina vorax Hirschfelder et al. 1993 Philodinavus paradoxus Madaliński 1961 Pleurata chalcicodis Jersabek et al. 2003 Pleurata tithasa Jersabek et al. 2003 Pleurata vernalis Jersabek et al. 2003

Pleuretra brycei Madaliński 1961

Pleuretra lineata Hirschfelder et al. 1993 Pleurotrocha robusta – Sphagnum bog Jersabek et al. 2003

Ploesoma lynceus Hingley 1993 Polyarthra euryptera Horkan 1931

Polyarthra minor* Hingley 1993 Polyarthra vulgaris Hingley 1993 Proales cognita* – S. cuspidatum Jersabek et al. 2003

Proales decipiens Horkan 1931; Hingley 1993 Proales doliaris – Sphagnum bog Hingley 1993;

Jersabek et al. 2003

Proales fallaciosa Hingley 1993 Proales latrunculus Hingley 1993 Proales micropus Hingley 1993 Proales minima Hingley 1993 Proales palimmeka* – submerged Jersabek et al. 2003

Proales sordida* Horkan 1931 Proales theodora Madaliński 1961

Proalinopsis caudatus Horkan 1931; Hingley 1993

Proalinopsis gracilis – Riccia fluitans Jersabek et al. 2003

Proalinopsis squamipes Hingley 1993;

Sphagnum ditch Jersabek et al. 2003 Pseudoploesoma formosum Jersabek et al. 2003

Ptygura brachiata – sessile on Sphagnum Hingley 1993

Ptygura cristata Edmondson 1940 Ptygura crystallina Horkan 1931 Ptygura elata Hingley 1993 Ptygura longicornis – sessile on Sphagnum Hingley 1993

Ptygura longipes – sessile on Sphagnum Hingley 1993

Ptygura melicerta Horkan 1931 Ptygura pilula – sessile on Sphagnum Hingley 1993

Ptygura rotifer – sessile on Sphagnum Hingley 1993

Ptygura velata – sessile on Sphagnum Hingley 1993

Resticula melandocus Hingley 1993 Resticula nyssa Hingley 1993 Rotaria haptica Hingley 1993 Rotaria macroceros Horkan 1931

Rotaria macrura Horkan 1931; Hingley 1993

Rotaria magna-calcarata Hingley 1993

Rotaria neptunoida Hingley 1993 Rotaria quadrioculata Hingley 1993

Rotaria rotatoria Horkan 1931; Madaliński 1961

Rotaria socialis Hingley 1993

Rotaria sordida Horkan 1931; Hirschfelder et al. 1993

Rotaria spicata Hingley 1993

Rotaria tardigrada Hingley 1993 Scaridium longicaudum Horkan 1931

Scepanotrocha rubra Horkan 1931; Hingley 1993

Squatinella bifurca* Jersabek et al. 2003

Squatinella longispinata Hingley 1993;

Sphagnum bog Jersabek et al. 2003

Squatinella microdactyla Hingley 1993 Squatinella mutica Hingley 1993

Squatinella rostrum (formerly S. mutica) Hingley 1993

Squatinella retrospina* – Sphagnum bog Jersabek et al. 2003

Squatinella tridentata Hingley 1993 Stephanoceros fimbriatus – sessile on Sphagnum Hingley 1993

Stephanoceros millsii Hingley 1993

Streptognatha lepta* Hingley 1993; Jersabek et al. 2003

Synchaeta pectinata Horkan 1931; Hingley 1993

Synchaeta tremula Horkan 1931 Taphrocampa annulosa Hingley 1993

Taphrocampa clavigera* Hingley 1993; Jersabek et al. 2003

Testudinella emarginula Hingley 1993 Testudinella epicopta – Sphagnum bog Jersabek et al. 2003

Testudinella incisa emarginula – Sphagnum bog

Jersabek et al. 2003

Trichocerca rattus Horkan 1931; Hingley 1993; Sphagnum bog Jersabek et al. 2003

Trichocerca rosea* in bog Hingley 1993; Jersabek et al. 2003 Testudinella patina Hingley 1993

Trichocerca rossae* Jersabek et al. 2003 Trichocerca rotundata* Jersabek et al. 2003

Tetrasiphon hydrocora* Norgrady 1980; Hingley 1993

Trichocerca brachyura Horkan 1931

Trichocerca scipi* Jersabek et al. 2003

Trichocerca similis Horkan 1931

Trichocerca bicristata Horkan 1931; Hingley 1993

Trichocerca cavia Hingley 1993

Trichocerca tenuior Horkan 1931; Jersabek et al. 2003 Trichocerca collaris Hingley 1993

Trichocerca elongata Hingley 1993

Trichocerca tigris Horkan 1931; Hingley 1993: Sphagnum and Riccia in pond Jersabek et al. 2003 Trichotria cornuta Jersabek et al. 2003 Trichocerca harveyensis – Fontinalis disticha

Jersabek et al. 2003

Trichotria pocillum Horkan 1931; Hingley 1993

Trichocerca junctipes Hingley 1993

Trichotria similis Jersabek et al. 2003 Trichocerca lata* Jersabek et al. 2003

Trichocerca longiseta Hingley 1993

Trichotria tetractis* Horkan 1931; Hingley 1993; Sphagnum in bog Jersabek et al. 2003 Trichotria tetractis caudatus Jersabek et al. 2003 Trichocerca ornata – Sphagnum bog Jersabek et al. 2003 Trichotria truncata Horkan 1931; Hingley 1993

Trichocerca parvula* Jersabek et al. 2003 Wierzejskiella elongata* Jersabek et al. 2003 Trichocerca platessa* Jersabek et al. 2003

Trichocerca porcellus Hingley 1993;

Fontinalis Jersabek et al. 2003

Wierzejskiella velox* Hingley 1993; Pejler & Bērziņš 1993;

Jersabek et al. 2003

CLASS BDELLOIDEA

This class of rotifers is exclusively parthenogenetic

(giving from unfertilized eggs), negating the need for males

to complete the life cycle. This group is comprised of ~460

species, only one of which is marine (Segers 2008). They

are distinguished from the Monogononta by the presence

of two ovaries (Monogononta have only one).

The bdelloids are known from freshwater and soil, and

are common on bryophytes. They have a retractable head

with a well-developed corona that is divided into two

parts. Movement includes both swimming and crawling,

but they seldom venture into the plankton (Fontaneto &

Ricci 2004). Crawling is similar to the movement of

inchworms, or some leeches. The name Bdelloidea is

derived from the Greek word meaning leeches, referring to

this method of movement.

Most of the bdelloids survive unfavorable periods,

particularly drought, by entering a type of dormancy known

as

anhydrobiosis (Gilbert 1974; Ricci 1987, 1998, 2001).

It is this ability, along with their parthenogenetic

reproduction (no male is needed) (Ricci 1992) that fosters

their cosmopolitan distribution (Fontaneto

et al.

2006b,

2007, 2008b). And this may also be the reason that Horkan

(1931), in his report on Irish rotifers, found only this group

on mosses other than those in bogs. Furthermore, no

Bdelloidea were present in the Irish bogs, on bog moss, or

in bog pools, suggesting they may need those dry periods.

Only one carnivorous bdelloid is known, and it is not

known from bryophytes. Rather, the bdelloids filter or

scrape or browse their diet of bacteria, one-celled algae,

yeast, or particulate organic matter (Ricci 1984).

Adinetidae

Ricci and Covino (2005) demonstrated various aspects

of anhydrobiosis in this family, using

Adineta ricciae

.

Rotifers that recovered from anhydrobiosis had similar

longevity and significantly higher fecundity than did the

hydrated controls. Lines of offspring produced after the

anhydrobiosis dormancy likewise had significantly higher

fecundity and longevity than controls from mothers of the

same age. The genus

Adineta

has many cryptic species, as

demonstrated by DNA and a diversity of narrow ecological

niches (Fontaneto

et al

. 2011).

Figure 2. Adineta barbata female, a species known to live

on Sphagnum subsecundum (Figure 3) and other mosses. Photo by Jersabek et al. 2003.

Figure 3. Sphagnum subsecundum. Photo by Michael Lüth.

Figure 4. Adineta gracilis, a species known from Sphagnum and other mosses. Photo by Jersabek et al. 2003.

Figure 5. Adineta vaga, a moss dweller that is 0.2-0.3 mm when extended. Photo by Jean-Marie Cavanihac at Micscape.

Habrotrochidae

Habrotrocha

species are common inhabitants among

Sphagnum

(Bateman 1987; Peterson

et al

. 1997; B

ł

edzki

& Ellison 1998). Habitats for

Habrotrocha

, in particular

H. rosa

, include pitcher plants (

Sarracenia purpurea

),

where they are a major food source for co-habiting

members of the Culicidae (mosquitoes) (Bateman 1987),

causing the mosquito population numbers to rise (B

ł

edzki

& Ellison 1998). The rotifers are an important source of N

and P in the bog/fen-dwelling pitcher plants.

Figure 6. Habrotrocha, a genus with many species that

occur on bryophytes. Photo by Proyecto Agua Water Project through Creative Commons.

Figure 7. Habrotrocha ampulla from among Sphagnum.

Photo by Jersabek et al. 2003.

Figure 8. Habrotrocha collaris female, a species known

from bryophytes. Photo by Jersabek et al. 2003.

Figure 9. Habrotrocha constricta female, a species known

from bryophytes. Photo by Jersabek et al. 2003.

Figure 10. Habrotrocha lata female, a species collected

from Sphagnum and other mosses. Photo by Jersabek et al.

2003.

Figure 11. Habrotrocha lata, a species collected from bryophytes in more than one location. Photo through EOL Creative Commons.

Philodinidae

The philodinids use their cilia or foot and proboscis

(Figure 26) to facilitate swimming (Hickernell 1917). At

high temperatures these rotifers engage in active

swimming, but in cold water they creep like a leech with

the cilia retracted. During feeding, they attach themselves

by the foot and use the cilia to direct food to the pharynx.

When drying occurs, the animal forms a ball and dries up.

The ball is formed by retracting both the head and the foot

into the trunk of the rotifer and losing all the water, pulling

the organs together and eliminating spaces. When they get

water again, they resume their normal shape in ten minutes

or less.

Figure 12. Dissotrocha aculeata female, a species known from Sphagnum and other mosses. Photo by Jersabek et al.

2003.

Figure 13. Dissotrocha macrostyla subsp. tuberculata

female, a species known from bryophytes in more than one location. Photo by Jersabek et al. 2003.

Figure 14. Macrotrachela ehrenbergii female, a species

known from Sphagnum. Photo by Jersabek et al. 2003.

Figure 15. Macrotrachela habita female, a species known

from Sphagnum and other mosses. Photo by Jersabek et al.

2003.

Figure 16. Macrotrachela multispinosa female, a species

known from "tree moss" and other mosses. Photo by Jersabek et al. 2003.

Figure 17. Macrotrachela multispinosa from among "tree moss." Photo by Jersabek et al. 2003.

Figure 18. Macrotrachela plicata, a species known from "tree moss" and other mosses. Photo by Jersabek et al. 2003.

Figure 19. Macrotrachela quadricornifera female, a species

known from Sphagnum and other mosses. Photo by Jersabek et al. 2003.

Figure 20. Macrotrachela sp., a genus with a number of species that live on Sphagnum. Photo by Walter Pfliegler.

Figure 21. Mniobia scarlatina from among "tree moss." Photo by Jersabek et al. 2003.

Figure 22. Philodina citrina female, a species known from Sphagnum bogs and "tree moss." Photo by Jersabek et al. 2003.

Figure 23. Philodina plena female, a species known from

Sphagnum. Photo by Jersabek et al. 2003.

Figure 24. Philodina roseola, a species known to inhabit bryophytes. Photo by Proyecto Agua Water Project through Creative Commons.

Figure 25. Philodina roseola females with eggs, a species known to inhabit bryophytes. Photo by Jersabek et al. 2003.

Figure 26. Rotaria macroceros, known from bog pools. The genus Rotaria is able to move among mosses and other substrata by creeping with its head and foot (van Egmond 1999). The foot (Figure 27) is sticky, enabling it to attach to a surface while it feeds (Dickson & Mercer 1966; Schmid-Araya 1998). The anterior cilia (Figure 28) make a current that directs the food toward the pharynx for ingestion. Note the proboscis. Photo from GLERL NOAA website.

Figure 27. Rotaria macrura from among Sphagnum and other mosses, showing fully extended foot. Photo by Jersabek et al. 2003.

Figure 28. Rotaria, showing the two wheels that direct the

food into the gullet. Photo by Yuuji Tsukii.

Figure 29. Rotaria rotatoria female, a species known from bryophytes in more than one location. Photo by Jersabek et al.

2003.

Figure 30. Rotaria, fully extended as it would be for its leech-like movement. This is a genus with several bryophyte -dwelling species that can move about the bryophytes in this manner. Photo by Wim van Egmond.

CLASS MONOGONONTA

This is the largest of the two classes of rotifers,

comprised of ~1570 species, ~1488 of which are free-living

in fresh water of limnoterrestrial habitats (Segers 2008). It

differs from the Bdelloidea in having two sexes and having

only one ovary. Nevertheless, asexual reproduction occurs

over and over until environmental conditions, often related

to crowding, trigger the reproduction to become sexual

(Welch 2008). At this time, the eggs of the amictic

(non-sexual) females hatch into mictic females that produce

their eggs by meiosis. The haploid eggs that are not

fertilized develop into much smaller males and fertilization

of a female by these males produces diploid eggs that

become resting eggs.

The

monogonont rotifers mostly eat small particles

and organisms by filtering them, some actually seize them,

and some are parasitic.

Order Collothecacea

Many members of this order are sessile (attached) and

some are colonial. These rotifers have a foot that lacks

toes, but they possess many foot glands that are used for

adhesion. The females are predominantly sessile, but

males and immature rotifers are free-living.. The rotary

apparatus surrounds a funnel-like invagination. Many are

surrounded with a jelly sheath.

Collothecidae

Many members of the Collothecidae are plant and

algal inhabitants.

Collotheca gracilipes

, a plant inhabitant,

is selective in its location on its substrate (Wallace &

Edmondson 1986). On plants such as

Elodea canadensis

, it

selected (98%) the lower (abaxial) surfaces of the leaves.

When given equal opportunities for four plant species, it

selected

Lemna minor

over

Elodea canadensis

, but in the

field more were found on

Elodea canadensis

, with densities

reaching more than six individuals per mm

_{. Light made a}

difference, with 91% of the rotifers selecting the adaxial

surface in continuous light, but showing no preference in

continuous darkness. Alpha amylase appears to be the

chemical that helps them to identify a plant substrate.

Those rotifers that were induced to settle on the abaxial

surface produced more eggs than those that were induced to

settle on the adaxial (upper) surface. It would be

interesting to see if these relationships persist on liverworts

like

Riccia fluitans

(Figure 31)

and

Ricciocarpos natans

.

But what would they do on mosses like

Fontinalis

?

Figure 31. Riccia fluitans, a substrate for Lecane flexilis

and other rotifer species, stranded here above water. Photo by Janice Glime.

The

Collothecidae provide us with evidence of

adaptive strategies embodied in reproduction. An

examination of 65 species of rotifers, including this family,

revealed that egg volume of rotifers increased as body

volume increased, but the relative size of eggs actually

decreased as body size increased (Wallace

et al

. 1998).

This means that smaller species, typical among planktonic

species, invest the most in egg production. The

Flosculariidae species are of intermediate size and their

relative investment in egg mass is likewise intermediate.

The

Collothecidae family has the largest species and the

lowest relative biomass of egg production among those

xamined by Wallace

et al

.

e

Figure 32. Collotheca, a common genus on Sphagnum.

Photo by Proyecto Agua Water Project through Creative Commons.

Figure 33. Collotheca sp., a common genus on Sphagnum.

Photo by Ed Purp through Micrographia.

Figure 34. Collotheca campanulata, a species that is known

as sessile on Sphagnum in bogs and occurs in bog pools. Photo by Jersabek et al. 2003.

Figure 35. Collotheca campanulata, a species that is known

as sessile on Sphagnum and occurs in bog pools. Photo by Yuuji Tsukii.

Figure 36. Collotheca catellina, a species known from bryophytes. Photo by Jersabek et al. 2003.

Figure 37. Collotheca catellina, a species known from bryophytes. Photo by Jersabek et al. 2003.

Figure 38. Collotheca coronetta, a species that occurs sessile on Sphagnum. Photo by Jersabek et al. 2003.

Figure 39. Collotheca crateriformis from among Sphagnum. Photo by Jersabek et al. 2003.

Figure 40. Collotheca crateriformis from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 41. Collotheca ornata, a species that lives in bogs and is sessile on Sphagnum. Photo by Jersabek et al. 2003.

Figure 42. Collotheca trilobata from among Sphagnum.

Photo by Jersabek et al. 2003.

Figure 43. Stephanoceros fimbriatus, a sessile species that

can occur ln Sphagnum. Photo by Wim van Egmond.

Figure 44. Stephanoceros fimbriatus female, a species that

occurs sessile on Sphagnum. Photo by Jersabek et al. 2003.

Figure 45. Stephanoceros millsii, a species known from bryophytes. Note the eggs. Photo by Jersabek et al. 2003.

Order Flosculariacea

Not only do the members of this order lack toes; some

of the planktonic species lack feet as well. Nevertheless,

they have multiple foot glands to secrete glue. The rotary

organ has a double ring of cilia that surrounds the anterior

of its lobe-like appendages. Species may be either

free-living or sessile and are suspension feeders.

Conochilidae

This family, or at least

Conochilus hippocrepis

(Figure 46, Figure 48), is sensitive to increasing predator

pressure from the copepod

Parabroteas sarsi

(Diéguez &

Balseiro 1998). As the predator increases in size and

begins to prey on the

Conochilus hippocrepis

, this rotifer

responds by increasing its colony size (Figure 47). The

only members of this family that seem to be known as

bryophyte associates are found among

Sphagnum

.

Figure 46. Conochilus hippocrepis subsp. unicornis female,

member of a genus known to associate with Sphagnum. The

species Conochilus hippocrepis is typically planktonic in both

ponds and large bodies of water, generally with a pH of 6.3-8.3

and temperature range of 6.4-15.4°C (de Manuel Barrabin 2000). Its colonies can reach 30-60 members that are joined in a gelatinous case. It eats detritus and bacteria (Pourriot 1977). Photo by Jersabek et al. 2003.

Figure 47. Conochilus sp. colony. This genus has species

that are sessile on Sphagnum. Photo by Wim van Egmond.

Figure 48. Conochilus hippocrepis female, member of a

genus known on Sphagnum. Photo by Jersabek et al. 2003.

Filiniidae

Only two members of the Filinidae seem to be known

from bryophytes:

Filinia longiseta

(Figure 49-Figure 50)

and

F. terminalis

(Figure 51). The latter lake species is

morphologically variable but seems to occupy a narrow and

well defined niche (Ruttner-Kolisko 1980). It prefers

temperatures below 12-15°C. At an oxygen content of less

than 2 mg L

_{, it can reach as many as 1000 individuals per}

liter. Not surprisingly, it is facultatively anaerobic. Its

food sources include bacteria that are chemosynthetic or

decompose plankton.

The members Filiniidae are highly variable and likely

comprise a number of microspecies (Ruttner-Kolisko

1989). This is at least in part due to the parthenogenetic

reproduction that can quickly lead to a clone of genetically

identical individuals in a founder population in a lake or

other habitat. This is furthermore complicated by the

absence of many good morphological characters by which

to distinguish species. In the

Filinia terminalis-longiseta

group, ecological properties differ and suggest the

existence of these microspecies, or perhaps species.

Figure 49. Filinia longiseta is known from bryophytes in England and Ireland. This is typically a cosmopolitan planktonic species of lakes, ponds, moorland waters, and even brackish water (de Manuel Barrabin 2000). It lives in a wide range of warm temperatures (7.7-26.2°C) and pH (6.3-9.9). It is a filter feeder on detritus, bacteria, and small algae like Chlorella in a size range of 10-12 µm (Pourriot 1965) and most likely competes for its food with members of the rotifer genus Conochilus. Photo by Jersabek

Figure 50. Filinia longiseta from bryophytes in a pond in Pennsylvania, USA. This species is also known from bog pools. Photo by Jersabek et al. 2003.

Figure 51. Filinia terminalis female, a cosmopolitan,

planktonic species known from bryophytes and Sphagnum bogs

(de Manuel Barrabin 2000). Its preferred conditions are mesotrophic to eutrophic in a pH range of 6.64-8.22. Its

temperature range is relatively wide: 7.3-22.8°C, although de Manuel Barrabin considers it to be a species of the cool hypolimnion. Photo by Jersabek et al. 2003.

Flosculariidae

In this family the male is small and free-swimming,

whereas the female lives in a tube and usually attaches by

its modified foot. Some of these females (

e.g. Ptygura

linguata

) live on the bladders of species of the bladderwort

Utricularia

. But, sadly for the rotifers, they also constitute

part of the diet of these same bladderworts (Mette

et al

.

2000). This habitat affords the rotifers a special aid in

getting food as it is sucked into the bladder. Bryophytes

can offer no such aid, and although the genera on

bryophytes are often the same because they are sessile,

species differ. As I read through account after account of

rotifer sampling, I couldn't help but wonder if more

attention should be given to the bryophyte habitat for

locating new rotifer species, especially for sessile groups

like this one.

Figure 52. Ptygura, a genus with a number of species known

to be sessile on Sphagnum, feeding among algae. Photo by

Micrographia.

Figure 53. Floscularia conifera female, a species that occurs sessile on Sphagnum and in bog pools. Photo by Jersabek

et al. 2003.

Figure 54. Ptygura, a common genus on bryophytes, showing its feeding cilia. Photo by Micrographia.

Figure 55. Ptygura sp with the green alga Spirogyra. Photo from Micrographia.

Figure 56. Ptygura brachiata female, known to be sessile on

Figure 57. Ptygura brachiata female, a species known to be

sessile on Sphagnum. Photo by Jersabek et al. 2003.

Figure 58. Ptygura crystallina female from the Pocono

Mountains, Pennsylvania, USA. This species has been collected with bryophytes and can occur in bogs. Photo by Jersabek et al.

2003.

Figure 59. Ptygura melicerta colony in a lake in Wisconsin,

USA. This species can occur among bryophytes and in bog pools. Photo by Jersabek et al. 2003.

Figure 60. Ptygura melicerta female from a lake in

Connecticut, USA. Here it is among Cyanobacteria; it can occur among bryophytes. Photo by Jersabek et al. 2003.

Figure 61. Ptygura melicerta colony in a lake in Wisconsin, USA. This species is known from bryophytes and bog pools. Photo by Jersabek et al. 2003.

Figure 62. Ptygura pilula female sessile on a Sphagnum

leaf; it also occurs in bog pools. Photo by Jersabek et al. 2003.

Figure 63. Ptygura rotifer female, a species known to occur

sessile on Sphagnum. Photo by Jersabek et al. 2003.

Hexarthriidae

In a study of a Turkish lake, Gülle

et al

. (2010) found

and disappeared from November through April. It was a

member of the Hexarthriidae,

Hexarthra fennica

that was

one of the dominant taxa – 51% of the zooplankton. The

rotifers were most dense at a depth of 5 m.

Figure 64. Hexarthra mira female from Mexico. This species is known from bryophytes and from bogs. Photo by Jersabek et al. 2003.

Figure 65. Hexarthra mira female from Mexico. This

species is known from bryophytes and from bogs. Photo by Jersabek et al. 2003.

Testudinellidae

The family Testudinellidae includes both salt water

and fresh water species. It is characterized by having

dorsal and ventral plates of the lorica that are completely

fused laterally. The body is greatly flattened

dorsi-ventrally. The foot is long and retractile (see Figure 68 and

Figure 73) with a tuft of cilia at its tip. These rotifers are

free-swimming, typically in the littoral zone, but members

of

Testudinella

may also occur on bryophytes and in

Sphagnum

pools as well as on other macrophytes. There

are three genera, but only

Testudinella

seems to be

represented on bryophytes.

Figure 66. Testudinella sp, a genus that occurs on bryophytes. Note the complete retraction of the foot. Photo by Wim van Egmond.

Figure 67. Testudinella clypeata, color modified. This

species is known from bryophytes and can occur in bogs. Photo by Leasi Francesca through EOL.

Figure 68. Testudinella epicopta from among Sphagnum.

Figure 69. Testudinella emarginula from a Sphagnum bog.

This cosmopolitan species lives on plant surfaces, although it occasionally occurs in the plankton (de Manuel Barrabin 2000). It is a cold water species (7.7-7.8°C) with a circumneutral pH

preference (pH 6..8-7.5) and wide alkalinity range. Photo by

Jersabek et al. 2003.

Figure 70. Testudinella incisa emarginula subsp emarginula from a Sphagnum bog. Photo by Jersabek et al.

2003.

Figure 71. Testudinella patina female. This is a planktonic species that likes small bodies of water where the aquatic plants are abundant (de Manuel Barrabin 2000). Bryophytes are among the aquatic plants in some associations where it has been found. The aquatic plant area provides it with its preferred foods of the green alga Chlorella and diatoms. It tolerates high salinity and lives in a pH range of 6.3-8.89. It enjoys a wide temperature

range of 9.5-24.3°C. Photo by Jersabek et al. 2003.

Figure 72. Testudinella patina; some members of this genus are Antarctic moss dwellers. Photo by Yuuji Tsukii.

Figure 73. Testudinella tridentata subsp dicella from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 74. Testudinella tridentata subsp dicella from among

Sphagnum. Photo by Jersabek et al. 2003.

Order Ploimida

This order has the most families. But are these species

ones likely to be on bryophytes? Wallace

et al

. (2008)

asked if "everything is everywhere?" They answered this

question in the Chihuahua Desert pools in Mexico. They

found that indeed the specialized, warm-water habitat of

the desert did not support "everything." The fauna was

dominated by families that are also common on

bryophytes: Brachionidae,

Lecanidae,

Lepadellidae,

and Notommatidae. Both habitats dry up.

Brachionidae

This is a family dominated by planktonic species and

was the family with the most species represented in

Spanish reservoirs (de Manuel Barrabin 2000), but a few

seem to spend time among bryophytes, perhaps as a place

to avoid predation, or just dropped there by moving water.

An interesting study by Stenson (1982) demonstrated,

however, that an experimental reduction of the fish

population led to an increase in larger rotifers and a

decrease in the smaller filter-feeding species such as

Keratella cochlearis

, a member of the Brachionidae.

Stenson attributed this to a change in competition for food

from rotifers such as

Polyarthra

(Figure 75).

Figure 75. Polyarthra major, a large rotifer that eats smaller

rotifers. Note the feather-like blades that are used like paddles in swimming. Photo by Wim van Egmond.

Feeding rates are inversely related to the density of

food organisms in

Keratella cochlearis

, as well as in

Polyarthra vulgaris

and

Polyarthra dolichoptera

(Bogdan

& Gilbert 1982).

Keratella

preferred

Chlamydomonas

to

all other foods offered, perhaps explaining its rarity among

osses, where

Chlamydomonas

also is rare.

m

Figure 76. Anuraeopsis fissa from a pond in Pennsylvania,

USA. This is a planktonic rotifer that has been found among bryophytes and in bog pools. It prefers warm water and a eutrophic habitat (Margalef 1955). It frequents small water bodies (de Manuel Barrabin 2000). Its food includes bacteria and detritus (Pourriot 1977) and it may become food for the rotifer

Asplanchna (Guiset 1977). Photo by Jersabek et al. 2003.

Figure 77. Anuraeopsis fissa from a pond in Pennsylvania,

USA. Photo by Jersabek et al. 2003.

Figure 78. Anuraeopsis fissa showing a single,

light-sensitive red eyespot and cilia. Photo from GLERL at plingfactory.

Figure 79. Brachionus urceolaris, a planktonic species that

is common in small, alkaline bodies of water (pH 7.25-9) (de

Manuel Barrabin 2000). It can occur in moving water and is relatively tolerant of high salinity. It is a cosmopolitan species with a wide temperature tolerance (7.35-24.3°C). Despite its alkaline preference, Hingley (1993) found it closely associated with Sphagnum in a bog. Photo from Smithsonian Institution.

Brachionus urceolaris

, and probably others, has a

survival trick against predation. The eggs survived

consumption by predators such as the cladoceran

Leptodora kindtii

without harm (Nagata

et al

. 2011). Often

the cladocerans would eject the eggs, and they typically

ejected the lorica while digesting the living contents. There

was a negative correlation between the portion of

unconsumed (ejected) eggs and the length of the predator.

Nevertheless, hatching success seemed to be independent

of the predator's body length. As many as 75% of the

undigested eggs hatched successfully.

Figure 80. Brachionus urceolaris, a planktonic species that

can occur in a Sphagnum bog. Photo by Michael Verolet.

Figure 81. Kellicottia longispina female, a central European

species known from bryophytes, is actually a planktonic species. Its long spines no doubt help to protect it from predation. It is active year-round as an inhabitant of oligotrophic lakes with a rather narrow pH range of 8.2-8.5, but as expected its temperature

range is broad (10.6-21.8°C) and it does not occur in small bodies of water (de Manuel Barrabin 2000). Its food is primarily chrysomonads and centric diatoms (Pourriot 1977). Photo by Jersabek et al. 2003.

Figure 82. Kellicottia longispina demonstrating spines that

may help in attaching it to bryophytes (Madaliński 1961). Photos GLERL at plingfactory.

Figure 83. Kellicottia longispina demonstrating spines that

may help in attaching it to bryophytes (Madaliński 1961). Photos GLERL at plingfactory.

Figure 84. Keratella mixta from among Sphagnum. Photo

by Jersabek et al. 2003.

Figure 85. Keratella quadrata female, a species known from bryophytes. This is also a cosmopolitan species that is active year-round (de Manuel Barrabin 2000). It is tolerant of mineralization and survives a wide pH range of 6.64-10.19. Its

temperature range is likewise wide (6.4-26.1°C), as expected for a perennial species. It has broad food preferences, including detritus, bacteria, and algae in the Chlorococcales, Volvocales, Euglenales, Chrysophyceae, and diatoms (Pourriot 1977). Photo by Jersabek et al. 2003.

Figure 86. Keratella quadrata female, a species known from bryophytes. Photo by Jersabek et al. 2003.

Figure 87. Keratella serrulata female. This is the only planktonic brachionid that is a specialist of acid water, particularly water from bogs with Sphagnum (Bērziņš & Pejler 1987). Its known pH is around 6.6 and temperature around

18.6°C (de Manuel Barrabin 2000). Photo by Jersabek et al.

2003.

Figure 88. Keratella serrulata feeds on algae in the

Chrysophyceae and Volvocales (Pourriot 1977). It lives in acid water, especially the outflow of Sphagnum bogs and poor fens.

Photo from GLERL at plingfactory.

Figure 89. Keratella serrulata showing rotary cilia. Photo

from GLERL at plingfactory.

Figure 90. Keratella serrulata female, a species known from

Sphagnum bogs and poor fen waters. Photo by Jersabek et al.

2003.

Dicranophoridae

The

Dicranophoridae are predators and are agile in

pursuing and capturing their prey (Pejler & B

ē

rzi

ņ

š 1993).

Unlike many rotifers, the Dicranophoridae are not

planktonic – other predatory rotifers exist there – and they

avoid the sediments where their prey organisms are not

sufficiently abundant. Unlike many rotifers, these have

been documented on two species of bryophytes through a

study of their substrata.

Albertia naidis

,

Aspelta angusta

,

A. aper

,

A. circinator

,

Dicranophorus forcipatus

,

D.

haueri

,

D. robusta europaeus

,

D. uncinatus

,

Encetrum

eurycephalum

,

E. fluviatile

,

E. lupus

,

and

E. mustela

were

all present on 1-10% of the 122 collections of

Fontinalis

.

Aspelta aper

,

A. circinator

,

Dicranophorus epicharus

,

D.

luetkeni

,

Encetrum arvicola

,

E. elongatum

,

E. incisum

,

E.

lupus

,

E. sutor

,

E. sutoroides

,

E. tyrphos

, and

Wierzejsklella velox

were all present on 1-10% of the 194

collections of

Sphagnum

. Both sets of bryophyte dwellers

occurred on a wide variety of plant substrata – none were

specific to bryophytes.

Whereas some families of rotifers are active

year-round, the Dicranophoridae are apparently sensitive to

warm weather. In a study of those members that live in the

interstitial spaces of a beach of the North Sea, the

Dicranophoridae can only be found in the cold seasons,

disappearing in mid-summer (Tzschaschel 1983).

Figure 91. Albertia naidis subsp intrusor from among Sphagnum and parasitic on Stylaria lacustris. This species is

also known from the aquatic moss Fontinalis. Photo by Jersabek

et al. 2003.

Figure 92. Trophus of Aspelta angusta from among mosses on rock. Photo by Jersabek et al. 2003.

Figure 93. Aspelta aper, a rotifer that occurs on both Fontinalis and Sphagnum species (Pejler & Bērziņš 1993).

Photo by Jersabek et al. 2003.

Figure 94. Aspelta beltista from among Sphagnum. Photo by Jersabek et al. 2003.

Figure 95. Aspelta circinator side view from among Sphagnum. This species is also known from bogs and Fontinalis. Photo by Jersabek et al. 2003.

Figure 96. Aspelta circinator from among Sphagnum.

Photo by Jersabek et al. 2003.

Figure 97. Aspelta chorista from among the moss Warnstorfia exannulata (formerly Drepanocladus exannulatus).

Photo by Jersabek et al. 2003.

Figure 98. Dicranophorus alcimus from among Sphagnum. Photo by Jersabek et al. 2003.

Figure 99. Dicranophorus artamus from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 100. Dicranophorus biastis from among Sphagnum. Photo by Jersabek et al. 2003.

Figure 101. Dicranophorus capucinus from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 102. Dicranophorus capucinus from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 103. Dicranophorus colastes from among Sphagnum. Photo by Jersabek et al. 2003.

Figure 104. Dicranophorus forcipatus, a rotifer found

among bryophytes in several studies. Upper Photo from the Smithsonian Institution, lower from GLERL NOAA.

Figure 105. Dicranophorus hercules capucinoides female, a species known from bryophytes. Photo by Jersabek et al. 2003.

Figure 106. Dicranophorus luetkeni female, a species

Figure 107. Dicranophorus luetkeni male, a species known

from Sphagnum. Photo by Jersabek et al. 2003.

Figure 108. Dicranophorus robustus female, a species

found with bryophytes in more than one location. Photo by Jersabek et al. 2003.

Figure 109. Dicranophorus robustus female, a species that is known to live among bryophytes and ingests members of the rotifer genus Lecane. Photo by Jersabek et al. 2003.

Figure 110. Dicranophorus rostratus female, a species

known from Sphagnum. Photo by Jersabek et al. 2003.

Figure 111. Dorria dalecarlica can occur on submerged moss in streams. Photo by Jersabek et al. 2003.

Figure 112. Encentrum felis female, a species known from bryophytes, including Sphagnum. Photo by Jersabek et al. 2003.

Figure 113. Encentrum felis from among Sphagnum.

Figure 114. Encentrum glaucum female, a species known

from bryophytes. Photo by Jersabek et al. 2003.

Figure 115. Trophus of Encentrum tobyhannaensis from

among Sphagnum. Often this is the only structure that can be

recognized in old collections. Photo by Jersabek et al. 2003.

Figure 116. Pedipartia gracilis from among Sphagnum subsecundum. Photo by Jersabek et al. 2003.

Figure 117. Streptognatha lepta female, a species known

from Sphagnum. Photo by Jersabek et al. 2003.

Figure 118. Streptognatha lepta female, a rotifer known to

associate with Sphagnum. Photo by Jersabek et al. 2003.

Figure 119. Wierzejskiella elongata from among

Sphagnum. Photo by Jersabek et al. 2003.

Figure 120. Wierzejskiella velox female, a species known

from Sphagnum in more that one location. Photo by Jersabek et al. 2003.

Epiphanidae

This family has rotifers that are usually planktonic, so

like most of the rotifers on bryophytes, it is likely that the

bryophyte is a temporary refuge. Many of the members of

this family are marine (Koste 1978; Fontaneto

et al

. 2006a,

2008a), where no bryophytes are known.

Figure 121. Cyrtonia tuba from a pond in Ohio, USA. This species has been collected from mosses. Photo by Jersabek et al.

2003.

Figure 122. Mikrocodides chlaena female from New Jersey,

USA. This species has been collected from mosses and from bog pools. Photos by Jersabek et al. 2003.