Principles of Systematics 1.

Introduction
Species taxonomy and Systematics: studying the diversity of life
classification Taxonomy and classification
Evolution
Biogeography
Conservation

By Luis Constantino

Keith Willmott
McGuire Center for Lepidoptera and Biodiversity
University of Florida
Gainesville, Florida, USA
Tropical Andean Butterfly Diversity Project -- Proyecto Diversidad de las Mariposas Andinas Tropicales

Lecture outline Activities of modern systematists

•Systematics, taxonomy and classification
• Delimitation and classification of species
•Linnaean taxonomy
• Reconstructing phylogeny
•Concept of taxon
• Higher classification
•Species and subspecies concepts
• Studying evolution of diversity
•Practical aspects of species description

Linnaean taxonomy The International Code of

• Standardised hierarchy (categories – taxonomic Zoological Nomenclature
rank): Kingdom, Phylum, Class, Order, Family,
Genus, Species Purpose: to promote clarity and stability
Principles:
• Standardised names for species and higher • Priority
groups • Flexibility: taxonomic changes permitted

• Species names consist of two parts – Adelpha Governs:

cocala; other groups a single name - Limenitidini • Publication
• Style and content of description
• Concept of taxon • Name priority and availability
• Type specimens
• Now governed by ICZN

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 Type specimens Species concepts
Holotype – the unique specimen
identified in original description Typological concept
Syntype – one of the type series Deviation from idealised
“type” requires new name

Neotype – the type specimen

designated after the original description
when no holotype or syntypes exist Adelpha ethelda Adelpha zalmona
Hewitson 1867 Hewitson 1871
Lectotype – a syntype chosen
after the original description to be the Parides phalaecus - syntype
unique type specimen Were early taxonomists
“typological” thinkers?
Paratype – part of the type series Main problem lack of
where a holotype is specified information on geographic
Allotype – a single designated variation
specimen from the type series of
opposite sex to the holotype Adelpha melona Adelpha deborah
Adelpha capucinus - neotype Hewitson 1847 Weeks 1901

Species concepts What is a subspecies? What is not a subspecies?

Biological species
concept
Actually or potentially
interbreeding populations
constitute a species;
Adelpha ethelda ethelda Adelpha ethelda zalmona trinomial nomenclature
Hewitson 1867 Hewitson 1871 Jordan, 1896
Consistently diagnosable
phenotype inhabiting a clearly
defined geographic region
• Inconsistently different
phenotypes
• Phenotypes occurring
sympatrically with other
phenotypes (polymorphism,
variation)
• Clinal variation (no discrete
variation)

Result
• great reduction in
number of “species”

Adelpha basiloides
Adelpha melona melona Adelpha melona deborah
Hewitson 1847 Weeks 1901 Mexico west Ecuador
Heliconius erato

Why have subspecies names?

Subspecies: problems and solutions
Problems:
1. Definition subjective (in contrast to species)
2. Variation usually clinal, inconsistent, minor
3. Subspecies are not incipient species sexmaculata lerdina lota

Response:
1. Also true for allopatric species and all higher taxa priscilla
2. Not true for many groups; don’t recognise ssp lerda
hermieri insolita
where true!
3. Not true for many groups, e.g. Heliconiinae,
Ithomiinae
ilderdina
gunilla camisea

2
 Why have subspecies names?
• information retrieval – taxonomic instability • heuristic taxonomy Biological species concept:
• improved understanding of taxon relationships • diversity patterns clarified
are two taxa species or subspecies?
Problems:
1. Hybridisation almost never observed in rare insects
2. Biological species concept cannot be applied to allopatric populations

sexmaculata insolita
lerda Solutions:
Oleria sexmaculata 1, 2. Assess reliability of characters to indicate taxonomic rank (species or
subspecies) based on variation in congeners

hermieri ilderdina priscilla Adelpha

Oleria ilerdina mesentina

camisea gunilla
lota lerdina
Oleria gunilla Adelpha iphiclus

Or… Cladistic, phenotypic etc

Biological species concept: species concept
special cases Species are monophyletic groups with at least one synapomorphy
Problems:
1. Some hybridisation observed among “true” sympatric species,
especially Heliconius
2. Some parapatric species hybridise rarely
Solutions: sexmaculata insolita
lerda
1. “Genotypic” species concept – maintenance of genetic integrity within Oleria sexmaculata
hybrid zone – are hybrids rarer than expected by chance?

hermieri ilderdina priscilla

Oleria ilerdina

Adelpha thesprotia Adelpha thesprotia x mesentina Adelpha mesentina

Jiggins, C. D., W. O. McMillan, W. Neukirchen, and J. Mallet. 1996. What can hybrid zones tell us about
speciation? The case of Heliconius erato and H. himera (Lepidoptera: Nymphalidae). Biol. J. Linn. Soc.
59:221-242. camisea gunilla
lota lerdina
Mallet, J. 1995. A species definition for the modern synthesis. Trends Ecol. Evol. 10(7):294-299 Oleria gunilla

Or… Cladistic, phenotypic etc So, what to do in practice?

species concept
Species are monophyletic groups with at least one synapomorphy • Use information on
range, habitat, ecology,
morphology
sexmaculata lerdina lota • Study wide range of
species and genera
A. seriphia aquillia
• Make decisions to be
priscilla
lerda
tested
hermieri insolita
• Obtain new information
and be flexible

ilderdina
gunilla camisea
A. seriphia therasia

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 Species description Systematics: Pachacutia baroni
Systematics Classification
• What is the genus? • Is the form already
• Is the form a species, described? mantura cleomella sp n
subspecies or variation? • Checklists, type specimens ?????????
Study sympatric and • Diagnosis
allopatric series • Description Genus
• What is the closest • Name formation, avoiding • New related to Brevioleria, Mcclungia
relative? homonymy • Well defined by 7 characters
• Is it sympatric or • Publication Godyris duillia Relatives
allopatric? • ICZN rules for species Allopatric to other species, but no sister
• If allopatric, sufficiently description species
distinct? Differences
Androconial and genitalic differences
Greta alphesiboea
Willmott, K.R., Lamas, G. 2007. A revision of Pachacutia, a new genus of rare Andean ithomiine butterflies
(Nymphalidae: Ithomiinae), with the description of two new species. Ann. Entomol. Soc. Am.

Colobura: species description

Classification: Colobura annulata
Is the form already described?
• 7 names already published for Colobura, from 1758 to 1942, based on
Atlas of Neotropical Lepidoptera checklist by Lamas
• dirce Linnaeus, bates Linnaeus, dirceoides Sepp – no type specimens so Diagnosis
use illustrations and descriptions, designate neotype
Description Sister species
sp. n., holotype identified, published in widely Male, female genitalia
circulated journal
Diagnosis Neotype dirce etc
Unique characters in larval colour pattern, adult wing pattern, male genitalia
Name formation, avoiding homonymy C. dirce C. annulata
Colobura annulata, from Latin annulatus meaning ‘ringed’, referring to
distinctive larval body rings – feminine adjective

Willmott, K.R., Constantino, L.M., Hall, J.P.W. 2001. A review of Colobura (Lepidoptera: Nymphalidae) C. dirce C. annulata
with comments on larval and adult ecology and description of a sibling species. Ann. Entomol. Soc. Am.
94(1): 185-196. Immature stages

Summary
• Butterflies one of the best studied groups
of organisms at species level
• New methods and philosophies greatly
changed butterfly taxonomy over the last
century, but now probably quite stable
• Still, even in well-known groups surprises
await…