Systematics and Phylogeny of benthic Copepods
Instructor:
Prof. Rony Huys, Department of Zoology, Natural History Museum, London.
e-mail: rjh@nhm.ac.uk
Israeli host: Prof. Bella Galil, Israel Oceanographic and Limnological Research, Haifa.
Dates: 06–10 November 2011.
Duration: 30 hours in total.
Location of the course: The National Collections of Natural History at Tel Aviv University
Objective of the course: to provide Israeli university students, postdoctoral fellows and other
professionals with a comprehensive and state-of-the-art introduction to the diversity, morphology,
systematics and biology of benthic, planktonic and symbiotic copepods, including training in
technical skills.
Prerequisites: As the course will be taught in English, applicants are expected to have a reasonable
command of the English language. Some basic knowledge of crustaceans and experience using a
dissecting (stereo) microscope are required.
Number of participants: maximum 15.
Grading: Oral examination, including species identification.
Background
No group of plants or animals on Earth exhibits the range of morphological diversity as seen among
the extant Crustacea. This structural disparity is best demonstrated by the Copepoda, which show an
immense vertical distribution – from the abyss to 5,500 m altitude, spanning three quarters of the
possible global vertical range on the planet. Copepods are aquatic microcrustaceans – the
microscopical relatives of the crabs and the shrimps and are often dubbed the “insects of the sea” –
usually ranging in size between 200 μm and 5 mm. They have colonized the biggest environment on
Earth – the massive 1,347 million km3 volume of water in the global oceans –, made the transition
from the sea to all freshwater habitats up to the Himalayan mountains, and have entered into
symbiotic relationships with virtually ever marine phylum, from sponges to chordates, including
mammals and reptiles. These three recurrent radiation events have made copepods the most abundant
multicellular group on the planet, even outnumbering the insects and the roundworms (nematodes),
both of which have had some claim to this position of pre-eminence. Their long evolutionary history
since their origin in the late Cambrian (500 Mya) has generated a plethora of life histories and body
shapes and modifications/reductions, to the extent that some copepod forms can no longer be
recognized as crustaceans without recourse to DNA sequence data. They underpin the world’s
freshwater and marine ecosystems, are sensitive bio-indicators of local and global climate change, key
ecosystem service providers, and parasites of economically important aquatic animals. Copepods
sustain the majority of world fisheries and through their roles as vectors of disease, also have a
number of direct and indirect effects on human health and quality of life.
The proposed 5-day training course will introduce participants to the biology of free-living and
symbiotic copepods, the high-level systematics and morphological characteristics of the group, the
main principles of zoological nomenclature and phylogenetic reconstruction, and the use of
identification keys. During the practicals participants will have the opportunity to collect copepods
from local coastal habitats and to learn dissection techniques and how microscopical observation can
lead to accurate identification. Practicals will focus on free-living marine benthic copepods and be
based on material provided by the tutor and/or from the collections made by IOLR, Haifa.
The course will consist of a combination of lectures, laboratory-based practical sessions and informal,
problem-solving discussions. Flexibility will be maintained to encourage the development of an
intense group dynamic through interactions between highly motivated participants and the course
tutor. A typical course day will consist of two 1.5-hour lectures alternated with practical sessions.
Course programme
Day 1
Introduction to the Copepoda. This session will provide a brief overview of copepod morphology and
diversity and introduce participants to the impressive range of body forms and limb types across the
Copepoda. Trainees will be introduced to the standardized terminology used in the description and
identification of free-living and symbiotic copepods.
Anatomy. An overview of the basic functional morphology of the reproductive, locomotory and
feeding systems will be presented, and the importance of the various sensory structures encountered in
copepods will be discussed.
Phylogeny. A brief synopsis of the current ordinal classification system will be presented, including
some recent developments in our phylogenetic understanding of the group.
Life cycles. Samples often have a high abundance of larval copepods which makes identification more
cumbersome since most of the taxonomy was established on the basis of adult morphology, and life
cycles are unknown for the vast majority of copepod species. Participants will be introduced to the
various life cycle stages and taught how to differentiate them.
Day 2
Taxonomic Techniques for the Study of Copepods. Topics will include procedures to extract material,
fix, preserve, stain and sort samples. The presentation will also demonstrate how to produce adequate
equipment to manipulate and dissect copepods and how to prepare temporary and semi-permanent
slides, focusing especially on the clearing, staining, mounting and dissecting techniques. The
information content of taxonomic illustrations will be discussed, and detailed information on
preparing line drawings and inking techniques will be provided.
Introduction to the marine benthic Copepoda. An overview will be given of the various families of
the order Harpacticoida – the most abundant copepod order in marine sediments – including notes on
their ecological preferences.
Laboratory Session I. Dissection and identification of tutor’s material. Using dichotomous and tabular
identification keys to identify copepods to family and genus level.
Day 3
Israeli harpacticoid fauna. A summary of our current knowledge on marine harpacticoid copepods
from the Levantine coast and Gulf of Elat. Participants will be introduced to the phenomenon of
Lessepsian migration.
Fieldwork. Brief sampling session on sandy and rocky shores in the Tel Aviv area.
Laboratory Session II. Dissection and identification of tutor’s and fresh material. Using dichotomous
and tabular identification keys to identify copepods to family, genus and species level.
Day 4
Zoological Nomenclature.Trainees will be introduced to the basic principles of nomenclature such as
availability, validity, synonymy, homonymy and typification.
Laboratory Session IV. Dissection and identification of tutor’s and fresh material. Using dichotomous
and tabular identification keys to identify copepods to family, genus and species level.
Day 5
Laboratory Session V. Dissection and identification of tutor’s and fresh material. Using dichotomous
and tabular identification keys.
Open Discussion and Problem-Solving Session
Oral examination.
Relevant literature
Boxshall, G.A. & S.H. Halsey. 2004. An Introduction to Copepod Diversity. The Ray Society,
London: i–xv, 1–966.
Hicks, G.R.F. & B.C. Coull. 1983. The ecology of marine meiobenthic harpacticoid copepods.
Oceanography and marine Biology. Annual Review 21: 67–175.
Huys, R. 2009. Unresolved cases of type fixation, synonymy and homonymy in harpacticoid copepod
nomenclature (Crustacea: Copepoda). Zootaxa 2183: 1–99.
Huys, R. & G.A. Boxshall. 1991. Copepod Evolution. The Ray Society, London: 1–468.
Huys, R., J.M. Gee, C.G. Moore & R. Hamond. 1996. Marine and brackish water harpacticoid
copepods. Part 1. In: R.S.K. Barnes & J.H. Crothers eds, Synopses of the British Fauna (New
Series), No. 51: i–viii, 1–352.
ICZN (International Commission on Zoological Nomenclature). 1999. International Code of
Zoological Nomenclature, Fourth Edition. The International Trust for Zoological Nomenclature,
London: i–xxx + 1–306. http://www.nhm.ac.uk/hosted-sites/iczn/code/
Lang, K. 1948. Monographie der Harpacticiden. Håkan Ohlsson, Lund. I: 1–896, figs 1–361; II:
897–1682, figs 362–607, maps 1–378.
Lang, K. 1965. Copepoda Harpacticoida from the Californian Pacific coast. Kunglige svenska
Vetenskapsakademiens Handlingar (4)10(2): 1–560.
Por, F.D. 1964. A study of Levantine and Pontic Harpacticoida (Crustacea, Copepoda). Zoologische
Verhandelingen, Leiden 64: 1–128.
Schminke, H.K. 2007. Entomology for the copepodologist. Journal of Plankton Research 29: i149–
i162.
Wells, J.B.J. 2007. An annotated checklist and keys to the species of Copepoda Harpacticoida
(Crustacea). Zootaxa 1568: 1–872.
Important websites
Copepod Web Portal by Lucio Pesce http://www.luciopesce.net/copepods/
World of Copepods http://www.nmnh.si.edu/iz/copepod
World Register of Marine Species (WoRMS) http://www.marinespecies.org/copepoda/
Taxonomic Techniques for Copepods by Janet W. Reid
http://invertebrates.si.edu/copepod/techniques.htm