DIET AND CRANIAL FUNCTIONAL MORPHOLOGY IN CANIDS: GEOMETRIC

MORPHOMETRICS AND THE COMPARATIVE STUDY OF ADAPTATION

Jesualdo A. 1
Fuentes-G. *, P. David 2
Polly , Joao 3
Muñoz-Durán , Krzysztof 4
Bartoszek , Jason Pienaar1

1The University of Alabama, 2Indiana University, 3Universidad Nacional de Colombia , 4Linköpings Universitet
INTRODUCTION The common prac ce of interpre ng shape trends obtained at boundaries OUTCOME
of the morphospace (e.g., Fig. 2C) is a useful exploratory tool but not very
Canid feeding preferences are remarkably variable (Fig. 1). appropriate for inferring adap ve pa erns, as it is inconsistent with the Preferred model is OU with four op ma (one for each diet). The op ma correspond to peaks in the morphospace which can be visualized (Fig. 6).
conceptual framework that inspired the formula on of the morphospace
(Fig. 3).
A B

Phenotypes

Fig. 3. Connec on between the adap ve landscape metaphor and the morphospace: (A) From Wright[4]: The
adap ve landscape as a polydimensional array of gene allele frequencies and a fitness axis. (B) From
Fig. 1. Main feeding preferences in extant canids: omnivory (e.g., Nyctereutes procyonoides), insec vory Simpson [5]: The adap ve landscape expanded to phenotypic spaces inspired the conceptualiza on of the
(e.g., Otocyon megalo s), carnivory (e.g., Vulpes lagopus), hypercarnivory (e.g., Canis lupus). morphospace; thus, op mum shapes are not necessarily expected at morphospace boundaries (as in Fig. 2C).

This feeding varia on should be reflected in skull shape, as this is the
main structure for obtaining and processing food in canids. The
morphospace offers a way to explore the skull-diet associa on (Fig. 2).
A B
AIM
Using the canid skull-diet associa on, we show how integra ng geometric
morphometrics with phylogene c compara ve methods offers an avenue to
model the morphospace in line with adap ve landscape thinking.

APPROACH

Using the morphogeometric data (Fig. 2) and a phylogeny[6], we apply[7] Fig. 6. Canid skull morphospace with shape op ma for each diet (filled dots colored as in Fig. 1). Consensus and ancestral reconstructed shapes also shown (empty dots).
C
variants of mul variate Brownian mo on (BM) and Ornstein-Uhlenbeck
(OU) models (Fig. 4). FINDINGS
x
x
x
x
x • Omnivory and insec vory cluster with consensus and ancestral reconstructed shape Stabilizing selec on
BM x
x
x
x OU
T x
x T x
x

dXT = ? * dWT x
x
x x
x
x
dXT = ? * dWT + a (? - XT) • Carnivory and hypercarnivory diverge from this cluster in opposite direc ons Disrup ve selec on
x x
x

µ ? • Omnivory undifferen ated (generalist); insec vory more differen ated with hypocarnivorous features (e.g., increased macera on surface).
Fig. 4. BM proper to model processes that generate varia on (e.g., neutral evolu on); OU proper to model
processes where varia on is constrained (e.g., selec on). • Diversifying trend reflects biomechanical trade-off between bi ng strength and occlussion speed: carnivory relies on slender, fast skulls (small
prey); hypercarnivory relies on robust, strong skulls (large prey).
BM and OU comprise various evolu onary modes that allow tes ng
adap ve hypotheses on the morphospace (Fig. 5). We compared various BM • These interpreta ons are less obvious when a connec on between the morphospace and the adap ve landscape is lacking (e.g., Fig. 2C).
(e.g, single or unequal rate matrices) and OU (e.g., number and structure of
adap ve peaks) models to iden fy which evolu onary mode explained CHALLENGES/FUTURE DIRECTIONS
canid skull shape be er.
• High risks of overparameteriza on: Mul variate compara ve models are
complex and morphogeometric data are highly dimensional.

Fig. 2. Canid skull morphospace from coordinate-based

[1] [2]
• Parameteriza on of compara ve models differs among implementa ons (e.g.,
data : (A) Landmarks placed on lateral view images (e.g.,
Cerdocyon thous). (B) Consensus shape based on en re
Landscape OUCH, mvSLOUCH, SURFACE, mvMORPH, l1ou, Phylogene cEM).
canid sample. (C) Morphospace generated by geometric
[3]
morphometric methods with shape trends shown at the
boundaries of each axis (filled dots represent species • In progress: Simula on study to assess effects of overparameteriza on, and to
grouped by convex hulls indica ve of diet; empty dot
represents consensus shown in B).
contrast the outputs generated by different implementa ons. Stay tuned…

ACKNOWLEDGMENTS
• Emília Mar ns for valuable support;
Elizabeth Housworth and Ellen
Process Neutral evolu on Randomly Stabilizing Func onal
fluctua ng selec on Ke erson for helpful discussions.
by dri selec on adapta on
• Art: Laura Fuentes.
• Design: Jorge Iván Atehortúa.
Model BM OU • This material is based on work
supported by fellowships to JAF-G
Fig. 5. BM useful to model landscapes that are effec vely flat; OU useful to model landscapes with discrete adap ve peaks (in the canid case, the number of peaks are informed by diet).
from Colciencias (Becas Caldas 497-
REFERENCES
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