Principles of Systematics 2.

Introduction
Higher classification • Names of people
• Identity and relationship
• Higher classes

Pelo negro

The Orkney Ba’ game

© Charles Tait
www.charles_tait.co.uk
Pelo rojo
Keith Willmott
McGuire Center for Lepidoptera and Biodiversity
University of Florida
Gainesville, Florida, USA Colombiano
Ingles
Tropical Andean Butterfly Diversity Project -- Proyecto Diversidad de las Mariposas Andinas Tropicales

Lecture outline Higher classification

Linnaean system
• Purpose of higher classification
• Hierarchy
– Linnaean nomenclature
• Governed (to some extent) by ICZN
– Rationale for higher classification
– Schools of systematics • Flexibility
• Higher classification – problems and • Stability
examples

Linnaean classification Goals of higher classification

• Categories or ranks
– Kingdom, Phylum, Class, Order, Family, Genus,
Species • Assist in identification
– A taxon is a named member of any category
• Continuing research on “Kingdom” relationships • Prediction of unknown traits
• Additional categories now also used: Linnaeus • Use in studies of evolution
described all butterflies in genus Papilio
• Superfamily, subfamily, tribe, subtribe • Data management and retrieval
Order Lepidoptera
Superfamily Papilion-oidea • Others?
Family Nymphal-idae
Subfamily Biblid-inae
Tribe Biblid-ini
Subtribe Eurytel-ina
Genus species Vila emilia

1
 History of higher classification New philosophies
• Assist identification
• Brief history of systematic thought:
• Reflect similarities: anatomy, biology
– Evolutionary systematics
• H.W. Bates, 1862: “Heliconidae”
• Predictive classifications based on phylogeny and
similarity
– Phenetics
• Minimise character distance; useful for identification
Heliconius numata Tithorea harmonia Lycorea halia – Cladistics
Heliconiinae Ithomiinae Danainae
• All taxa are monophyletic
• Changing philosophies with Darwin’s theory of
evolution in 1859: desire to reflect phylogeny • Third dominant but in practice elements of
• Phylogenetics and higher classification linked all three are used

Phylogenetics and higher classification Phenotypic vs genealogical similarity

Monophyly, paraphyly and polyphyly C D E

Monophyletic group = clade Phenetics, evolutionary systematics:

Family 1: A, B
Monophyletic group Paraphyletic group Polyphyletic group Subfamily I: A
= clade, includes ancestor Includes the common Doesn’t include common Subfamily II: B
and all descendants ancestor and only some ancestor or all descendants Family 2: C, D, E
of the descendants Subfamily III: C, D, E

O A B C D E O A B C D E O A B C D E Cladistics:
Family 1: A
Subfamily I: A
B Family 2: B, C, D, E
A Subfamily II: B
Subfamily III: C, D, E

Valid taxon Invalid taxon Synapomorphy

Application of cladistic methods to General lessons

neotropical butterfly classification • Classification from single character
W&F – Some relationships mistaken: Aeria
Fox

Tithorea

Aeria

Example:
Oleria
Ithomiinae

2
 General lessons General lessons
• New information • Application of cladistic
– Autapomorphic taxa show changes in philosophy Equally parsimonious

classification: Methona – Primitive groups often

paraphyletic: Tithoreini
– Diverse, homogeneous taxa often turn out to
• Members often
be polyphyletic and require reclassification: “conservative”
e.g. Hypoleria • Difficulty in finding
synapomorphies
– Primitive taxa often
mis-classified:
Velamysta
“Hypoleria” sarepta

Methona grandior

Description of higher taxa Information for new higher taxa

• study all taxa in group • wing pattern of representative males and females
• study all other plausibly related taxa and close relatives illustrating synapomorphies
• study as many characters as possible, • genitalia of representative males and females and
including morphological and molecular where
possible close relatives illustrating synapomorphies
• verify that no applicable names exist • other morphological structures
• description: includes interspecific variation and • molecular sequences that are synapomorphies
characters unique to genus
• diagnosis: includes synapomorphies verified
by cladistic analysis including close relatives
• type must be specified: should be well known Morphology of
“Pachacutia, a new
• etymology: ensure name formed correctly; genus of Ithomiinae
(Willmott & Lamas,
check for homonyms in press)

Justification for description of new higher taxa Example: classification

in Riodinidae
• group is monophyletic • albinus, glaphyra need a new genus
• group is heuristic (ie included taxa likely to share features) • odites transferred from Synargis to Juditha
• group is readily identified (preferably via visible features) - Why not new genus? Shares unique
synapomorphy with remaining
• minimise changes in current usage (stability) congeners and not ecologically distinct
• sister taxon also monophyletic Unique androconial setal pad
between valva and pedicel in
Juditha – from Hall & Harvey
(2001)
EXISTING PROPOSED
Genus A Genus A New genus • several “Audre” species moved to Lemonias
• Uraneis synonymised with Thisbe – though From Hall & Harvey (2001)
D E D E distinct, make Thisbe paraphyletic
O O
Describer responsible for
resolving relationships in
A B C A B C Genus A irenea lycorias ucubis
Hall, J.P.W., Harvey, D.J. 2001. A phylogenetic analysis of the neotropical riodinid butterfly genera Juditha,
Lemonias, Thisbe and Uraneis, with a revision of Juditha (Lepidoptera: Riodinida: Nymphidiini). Syst. Ent.
26:453-490.

3
 Summary
• despite long history of taxonomic study, much
higher taxonomic work remains to be done
• cladistic principles and computers have
revolutionised butterfly taxonomy
• new character sources also important, including
morphology (immature stages, ultrastructure), but
especially molecular sequences
• real promise of reaching stable higher
classification for all butterflies within 10-15 years