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Geometric morphometrics and cladistics: Testing evolutionary relationships in mega- and microbats


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Traditionally, morphometric data have consisted of distances, angles, or ratios, and have been considered inappropriate for cladistic analyses. Recently, geometric morphometrics, based on homologous landmark point-coordinates, has provided a number of advantages over traditional morphometric data and methods, including the possibility that phylogenetically informative characters and character-states may be extracted and used in cladistic analyses. Using two data sets of 3-dimensional point coordinates collected from skulls of bats, we empirically evaluate this possibility. Partial warps were extracted from the point-coordinate matrix, and these were then re-coded by gap-coding, for use in the cladistic analyses. In the case of samples from Eidolon helvum populations (two mainland localities and four islands in the Gulf of Guinea), analyzing males and females separately, our analyses based on these data were unable to detect consistent phylogeographic patterns among the populations. In the case of samples from plecotine bat species, these analyses produced a consensus cladogram showing considerable concordance with an earlier cladistic analysis by us of this group. In both cases, our results reflect those of earlier studies (based on both morphologic and genetic data), suggesting that the data and analytic techniques described herein may have interesting utility in cladistic analyses.
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It has been argued that continuous-state
characters should be excluded from cladis-
tic analysis mainly due to two reasons: (1)
these data are inappropriate (not phyloge-
netically informative), and (2) the methods
for their conversion into codes are arbitrary.
Pimentel and Riggins (1987) stated that
only character states derived from charac-
ters showing discrete variation can provide
phylogenetic information, whereas continu-
ous characters are transformational homolo-
gies not subject to test and, therefore invalid
for phylogenetic analysis. On the other
hand, more recently Rae (1998: 221) con-
cluded that “metric data [...] fulfill the sole
criterion for inclusion in phylogenetic
analysis, the presence of homologous char-
acter states, and thus cannot be excluded as
a class of data.” This position is also sup-
ported by Thiele (1993), who noted that
data scored for cladistic analyses may be
quantitative or qualitative, continuous or
discrete, and show overlapping or non-over-
lapping values between taxa.
Traditionally, morphometric research has
relied on statistical analyses of distances,
Acta Chiropterologica, 7(1): 39–49, 2005
PL ISSN 1508-1109 © Museum and Institute of Zoology PAS
Geometric morphometrics and cladistics: testing evolutionary
relationships in mega- and microbats
1Museum and Institute of Zoology, Polish Academy of Sciences, 00-679 Warszawa, Poland
2Estación Biológica de Doñana (CSIC), Avda. Mª Luisa s/n, Sevilla 41013, Spain
3Department of Biological Sciences, Texas Tech University, Lubbock, TX 79409, USA
Traditionally, morphometric data have consisted of distances, angles, or ratios, and have been considered
inappropriate for cladistic analyses. Recently, geometric morphometrics, based on homologous landmark point-
coordinates, has provided a number of advantages over traditional morphometric data and methods, including
the possibility that phylogenetically informative characters and character-states may be extracted and used in
cladistic analyses. Using two data sets of 3-dimensional point coordinates collected from skulls of bats, we
empirically evaluate this possibility. Partial warps were extracted from the point-coordinate matrix, and these
were then re-coded by gap-coding, for use in the cladistic analyses. In the case of samples from Eidolon helvum
populations (two mainland localities and four islands in the Gulf of Guinea), analyzing males and females
separately, our analyses based on these data were unable to detect consistent phylogeographic patterns among
the populations. In the case of samples from plecotine bat species, these analyses produced a consensus
cladogram showing considerable concordance with an earlier cladistic analysis by us of this group. In both
cases, our results reflect those of earlier studies (based on both morphologic and genetic data), suggesting that
the data and analytic techniques described herein may have interesting utility in cladistic analyses.
Key words: geometric morphometrics, partial warps, gap-coding, phylogeny, Microchiroptera, Megachiroptera
angles or ratios to study quantitative vari-
ation among taxa. Recently, geometric
techniques have been developed for the di-
rect analysis of coordinates of homologous
landmarks. This geometric morphometrics
methodology has several advantages over
the traditional morphometrics (sensu Mar-
cus, 1993), e.g., by partitioning size from
shape for separate analyses and by provid-
ing a graphic way of locating and compar-
ing variability in different components of
shape among studied groups (e.g., Rohlf
and Marcus, 1993). Recently a number of
arguments have been made in favor of re-
garding certain morphometric variables as
putatively homologous characters and in-
cluding them, sometimes along with other
non-morphometric variables, in parsimony-
based cladistic analyses. Zelditch et al.
(1995; see also Fink and Zelditch, 1995 and
Swiderski et al., 1998; but cf. Adams and
Rosenberg, 1995; Naylor, 1996; and Mon-
teiro, 2000) have proposed the use of partial
warp-based traits in phylogenetic analysis.
These traits are derived from the shapes of
objects under study, as defined by the se-
lected number of x, y(and zin 3-dimension-
al analysis) coordinates of homologous
landmarks. The shapes are fitted to the ref-
erences by stretching/compressing and
shearing until complete identity of their
landmark configurations is achieved. Eigen-
vectors of the resulting bending-energy ma-
trix are defined as new shape variables,
principal warps which yield another shape
space with the origin defined by the refer-
ence. Projections of the shapes being com-
pared onto principal warps yield partial
warps, which warps are analogous to factor
analysis projections, with the eigenvectors
(principal warps) derived from a description
of non-uniform differences between ob-
served forms and some reference form
(O’Higgins, 2000). Principal warps, togeth-
er with the uniform component, also supply
a basis for the space in which we compute
relative warps, which are the same as prin-
cipal components (in particular, they are
mutually orthogonal). Both partial and rela-
tive warps can be used in many multivariate
statistical analyses as quantitative shape
variables. MacLeod (2002) illustrated some
of the weaknesses of partial warps using
empirical and simulated examples. Unfortu-
nately, the empirical trilobite example used
by this author is not a good test case be-
cause, as noted by Wagner (2000), trilobite
matrices are characterized by poor resolu-
tion of states.
The aim of our study was to re-evaluate
the usefulness of methods applying partial
warp analysis based on 3-dimensional in-
formation in recovering phylogenetical-
ly informative characters. This was tested
by producing trees derived from classical
(linear) parsimony of re-coded data, and
squared-change parsimony and continu-
ous maximum likelihood of continuous
characters, and comparing them with
a presumably known and well-established
phylogeny of two groups of bats belong-
ing to Megachiroptera and Microchiro-
For testing interspecific relationships among fruit
bats four taxa occurring in western Africa and with
well-understood phylogenetic relationships were se-
lected: Eidolon helvum, Rousettus aegyptiacus, Myo-
nycteris torquata and M. brachycephala. In addition,
the first species was also used in evaluation of evolu-
tionary ties among its populations in the islands of the
Gulf of Guinea. Eidolon helvum is the second largest
fruit bat in Africa, with unique morphological (An-
dersen, 1912), ecological (Thomas, 1983), and repro-
ductive (Bernard and Cumming, 1997) characteris-
tics. Juste et al. (2000) reported that of four island
populations examined (plus two mainland popula-
tions), the population from Annobón, the smallest and
farthest, shows remarkable morphological and genet-
ic differentiation, whereas the rest are similar phenet-
ically and with low genetic distances among them.
40 W. Bogdanowicz, J. Juste, R. D. Owen, and A. Sztencel
A set of 20 homologous cranial landmarks (see
Bookstein, 1991) was defined on bone sutures, foram-
ina, and inflection points along the edges of cranial
structures (Fig. 1) on the ventral view of skulls of the
four species of fruit bats (Appendix). Dental land-
marks were set on the bone (at the edge of the alveoli)
to avoid variation due to differential tooth-wear. To
facilitate repeatability, each landmark was gently
marked in pencil on the surface of the bone, under
20x magnification, before being recorded. Three-di-
mensional coordinates of landmarks were digitized
using a 3-D Reflex Microscope (Reflex Measurement
Ltd., Butleigh, Somerset BA6 8SP, UK). This is a
highly precise, non-contact instrument that uses a
small light spot to digitize coordinates in any position
within a magnified field. The microscope was period-
ically re-calibrated to ensure a linear scale error of
less than 30 µm over 100 mm in the x-axis. Land-
marks were collected under a 20× magnification lens
from four different aspects, thus the skull was rotated
three times to attain the complete data collection.
Amount of error due to the new digitizing of the four
reference landmarks was evaluated after each shift,
and the whole transformation was discarded, and the
digitizing process restarted, if the greatest error of any
of the four reference points was larger than 0.1 mm in
any direction. All these landmarks were recorded by
a single individual, and were taken without reference
to prior values.
The same instrument was applied to gather 19
three-dimensional coordinates from the dorsal side of
the skulls of microbats, including members of the
tribe Plecotini sensu stricto (genera Corynorhinus,
Plecotus, Barbastella, Euderma, Idionycteris) and
some other taxa (Otonycteris hemprichi, Antrozous
pallidus, and Eptesicus fuscus) within the subfamily
Vespertilioninae, plus Myotis lucifigus of the subfam-
ily Myotinae (sensu Hoofer and Van Den Bussche,
2003), which was used as an outgroup (see Appen-
dix). The plecotine bats represent the only supra-
generic group within Chiroptera that is Holarctic in
distribution, and there is considerable morphologic
and karyotypic evidence supporting their monophyly
(e.g., Frost and Timm, 1992; Tumlison and Douglas,
1992; Bogdanowicz et al., 1998; see also Juste et al.,
2004). Also in this case all landmarks were recorded
by a single individual.
Geometric coordinates were checked and visual-
ized using Morphologika (O’Higgins and Jones,
1998). The average location of each landmark (if
necessary) was obtained through a non-documented
Geometric morphometrics and cladistics in mega- and microbats 41
FIG. 1. Location of landmarks on the skulls of Megachiroptera (left) and Microchiroptera (right)
option in the program allowing to save the coordi-
nates of the warped mean (P. O’Higgins, in litt.). The
NTSYS-pc ver. 2.11T (Rohlf, 2000) package was
used to calculate partial warps. The uniform compo-
nent was estimated by sweeping the partial warps
from the projections of the aligned coordinates into
the tangent space and then using SVD to extract the
non-singular dimensions (Rohlf and Bookstein,
2003). Partial warps (plus uniform components
scores) were re-coded using the gap weighting
method of Thiele (1993). A new character state Xnew
can be calculated with the following formula:
Xnew = n* [(x - min) / (max - min)],
where max and min are the maximum and minimum
mean value of the character across all species, x is the
mean value of the current taxon and nis the number
of allowed character states.
Phylogenetic analyses were performed with
PAST (Hammer et al., 2001) and PHYLIP (Felsen-
stein, 2004b). The first taxon was always (but inten-
tionally) treated as the outgroup. In the case of fruit
bats the branch-and-bound algorithm was applied.
For the Plecotini project we used heuristic search,
with the subtree pruning and regrafting option. This
algorithm is similar to the nearest neighbour inter-
change but with a more elaborate branch swapping
scheme. The character optimization was based on the
Wagner criterion, assuming that characters are re-
versible and ordered, meaning that 0- > 2 costs more
than 0- > 1, but has the same cost as 2- > 0.
Fruit Bats
In the cladogram (Fig. 2A) representing
interspecific relationships among these
megabats Rousettus aegyptiacus figures as
next most basal to the outgroup, followed
by the Myonycteris clade, which contains
M. brachycephala and M. torquata.
In the cladograms representing intraspe-
cific relationships of the E. helvum popu-
lations, Rousettus is seen as basal, with
a clade of the two Myonycteris species
the next most basal (Fig. 2B and 2C). The
relationships among the Eidolon popu-
lations are figured differently, depend-
ing on whether based on the male (tree of
936 steps, ensemble consistency index, CI
= 0.76) or female shortest tree (1,002 steps,
CI = 0.79) data. In the cladogram based on
42 W. Bogdanowicz, J. Juste, R. D. Owen, and A. Sztencel
FIG. 2. Interspecific relationships (A) among four
species of fruit bats derived from the maximum
parsimony of re-coded partial warp scores (strict
consensus tree). Interpopulation affinities on the basis
of two equally long cladograms in E. helvum are also
shown (B, XX; C, YY). Numbers below branches
indicate the bootstrap support values (percentage) for
the same nodes selected in the topologies obtained
under the evolutionary model of maximum
parsimony (after 100 iterations and 500 reorderings).
Please note that in each case only one possible
orientation to the reference is taken into account
female shape characteristics (Fig. 2B), pop-
ulations from Annobón and Principe are
sister taxa, forming a sister group to the
remaining four populations studied. Within
these four, Río Muni is basal, followed
by Bioko, which is sister to the Cameroon/
São Tome clade. In the male-based clado-
gram (Fig. 2C), Principe is again in the
most basal group, but is sister to Río Muni.
Together these two populations are sister
to the remaining four, among which São
Tome is most basal, followed by Bioko,
which is sister to Cameroon and Anno-
bón. Thus, there is little or no concordance
to be found between the interpopula-
tional cladograms based on the female
and male shape characteristics, and neither
represents a recognizable geographic pat-
The three shortest trees were 1,110 steps
long and had an ensemble CI of 0.70. The
strict consensus (Fig. 3A) reflects a high de-
gree of concordance among the three clado-
grams, which are in agreement concerning
the placement of the non-plecotine taxa
Geometric morphometrics and cladistics in mega- and microbats 43
FIG. 3. Relationships among bats belonging to the tribe Plecotini: A — consensus of three most parsimonious
trees from the analysis of re-coded partial warps; B — the most parsimonious tree based on morphological and
karyotypic evidence (Bogdanowicz et al., 1998); C — extracted super-tree of Plecotini (Jones et al., 2002);
D — cladogram for Plecotini based on ca. 2.6 kb pairs of mitochondrial DNA sequence (after Hoofer and Van
Den Bussche (2003), but with only relationships supported strongly by either or both Bayesian and Parsimony
analyses being depicted
44 W. Bogdanowicz, J. Juste, R. D. Owen, and A. Sztencel
Otonycteris,Eptesicus, and Antrozous (out-
groups treated analytically as ingroups).
The ‘correct’ arrangement of these taxa sug-
gests that our characters: (a) carry phyloge-
netic information at this taxonomic level,
and (b) are correctly polarized.
Within the plecotine taxa, the genus Ple-
cotus is sister to all other taxa, with P. aus-
triacus and P. kolombatovici sisters, with
P. auritus a sister to that clade. The remain-
ing taxa are figured as an unresolved poly-
tomy among the genera Euderma, Idiony-
cteris, Corynorhinus (2 species), and Bar-
bastella (2 species). (The majority consen-
sus of the three trees agrees with this, but
shows Euderma and Idionycteris as sister
species, and an unresolved trichotomy
among this clade, Corynorhinus, and Bar-
Interspecific and Intrapopulation Rela-
tionships of Fruit Bats
The resulting phylogenetic hypothesis at
the species level was consistent with the
previously established arrangements of
fruitbats based jointly on sequences derived
from the mitochondrial cytochrome b and
16S rRNA genes of a wide representation of
Megachiroptera (Álvarez et al., 1999; Juste
et al., 1999; see also Juste et al., 1997), de-
spite outstanding differences in the evolu-
tionary relationships among fruitbats sug-
gested by molecular and ’classical‘ morpho-
logical data (Springer et al., 1995; Kirsch
and Lapointe 1997). On the other hand, it is
not surprising because only a four-taxon hy-
pothesis was tested using re-coded partial
warps, including two taxa (M. brachycepha-
la and M. torquata) belonging to one genus
(Romagnoli and Springer, 2000).
The situation is much more complicated
in the case of evolutionary affinities of
a single species — Eidolon helvum — on
the islands of the Gulf of Guinea (Central
Africa). An Eidolon ancestor probably
reached mainland Africa independently
from other fruitbats, and this colonization
likely took place far earlier than the Late-
Pliocene date (3 Myr) of its only fossil
(Howell and Coppens, 1974) and maybe
even earlier than the other African coloniza-
tions (Juste et al., 1999). A recent phyloge-
netic study suggests that Eidolon’s origin
may be closer to the typically Asian Ptero-
pus group than to any extant African fruit
bat (Juste et al., 1999).
In ‘classical’ multivariate morphology,
the populations of E. helvum from the is-
lands of Bioko, Príncipe, and São Tomé
do not show significant phenetic differenti-
ation, although a trend towards a reduction
of size is found in the latter two islands (Fig.
4). In terms of allozyme variation, the low
genetic distances among these populations,
as well as their values of Wright’s fixation
indexes, suggest that gene flow has ham-
pered differentiation on these islands (Juste
et al., 1999). Only the fourth insular pop-
ulation, from Annobón, was characterized
by such remarkable morphological and ge-
netic differentiation that it has been accord-
ed the status of a separate subspecies:
E. helvum annobonensis. Although we
might expect better or more consistent reso-
lution among these island populations based
on re-coded partial warps, it appears that
resolution of our approach mirrors that of
more traditional approaches, and is unable
to discern phylogeographic patterns among
the Eidolon populations. Still, the partial
warp analysis is able to tell apart the mor-
phological differences specific of the fe-
males of the population from Annobón.
Interspecific Relationships within Plecotini
A comparison of our strict consensus
tree with the pertinent portion of the
’supertree‘ of Jones et al. (2002) is of
considerable interest (see Fig. 3A and 3C),
in that if the rooting is disregarded, the
Geometric morphometrics and cladistics in mega- and microbats 45
FIG. 4. The geographic situation in the Gulf of Guinea, Central West Africa (based on Juste et al., 2000). The
neighbour-joining topologies based on selected morphological characters, by sex, built on Mahalanobis D2
distances between populations of E. helvum from the Gulf of Guinea, according to Juste et al. (2000), are also
topologies are concordant (albeit with dif-
fering resolution). Considering only the ple-
cotine taxa, and configuring the topology of
Jones et al. (2002) to reflect a primary di-
chotomy of Plecotus as sister to all other
plecotine taxa (as in our consensus tree),
their tree would figure Corynhorhinus as
sister to a clade containing Barbastella and
Euderma/Idionycteris. Although our con-
sensus tree figures these last three clades as
an unresolved trichotomy, the topologies
are concordant. Thus, the difference be-
tween our phylogenetic hypothesis for the
plecotines, and that of Jones et al. (2002), is
one of polarization, or more precisely, the
ordering of character-state transformation
series. In fact our consensus tree is more
likely correct than Jones et al. (2002), be-
cause it is very difficult to postulate that
the Euderma/Idionycteris clade is basal to
a clade containing both Plecotus (Old
World) and Corynorhinus (New World). If
we assume that the Plecotini originated in
the Old World, their phylogeny requires ei-
ther: (a) two dispersals from Old to New
World, or (b) one dispersal from Old to New
World, followed by a dispersal from New to
Old World; whereas our phylogeny could be
resolved to require only one dispersal from
Old to New World (if we allow Euderma
and Idionycteris to be sisters, as in our strict
consensus tree, and further resolve the tri-
chotomy to allow the Euderma/Idionycteris
clade to be sister to Corynorhinus), which is
reasonable on biogeographic grounds.
We also note that the pertinent portion of
the tree presented by Hoofer and Van Den
Bussche (2003) is concordant with our con-
sensus tree, though their data were unable to
provide much resolution at this level, or
even demonstrate that the Plecotini are
monophyletic. They did agree, however, on
the sister-group relationship between Eu-
derma and Idionycteris.
Geometric Morphometrics, Partial Warps
and Phylogeny
In the present study, in the case of
samples from plecotine bat species, these
analyses produced a reasonable consensus
cladogram showing considerable concor-
dance with an earlier cladistic analysis by us
of this group, and our results reflect those of
earlier studies (based on both morphologic
and genetic data; e.g., Frost and Timm,
1992, and Bogdanowicz et al., 1998). The
most fundamental problem in using tradi-
tional morphometric shape variables in a
standard cladistic analysis is the fact that it
requires the use of Manhattan distances (it
is implied by the use of linear parsimony).
Geometric morphometrics yields variables
corresponding to an arbitrary rotation of
shape space (depending in part on the refer-
ence configuration orientation). This arbi-
trariness does not matter in morphometrics
because the multivariate methods used in
morphometrics are invariant to rotation.
Manhattan distances are not and thus one
must select a particular rotation as being es-
pecially meaningful. Gap coding applied
one variable at a time is also a problem be-
cause the results depend on the arbitrary ro-
tation of the space (e.g., a different orienta-
tion of the reference configuration will yield
different shape variables that will be coded
differently by gap coding) (reviewed in
MacLeod and Forey, 2002; see also Felsen-
stein, 2004a). The squared change parsimo-
ny does not have this problem so that would
be quite compatible with morphometric data
since the solution is invariant to rotation of
the data. Continuous maximum likelihood
methods are also compatible with morpho-
metric data (F. J. Rohlf, in litt.). Neverthe-
less, at least as far as complex morphologi-
cal structures are concerned, such models
have probably no biological meaning. Mo-
reover, despite forcing continuous data into
integer codes only the classical, linear parsi-
mony of re-coded partials warps revealed
considerable logical phylogenetic configu-
ration for a 13-taxon example of Plecotini,
contrary to the methods utilizing continuous
data. It appears that a period of active ex-
perimentation with these methods is now
needed to further explore their appropriate-
ness and compatibility. An approach worth
investigating in this context is presented by
Bookstein (2000, 2002).
The following institutions are acknowledged for
allowing us to examine specimens under their care
and their curators and/or collection managers for fa-
cilitating the loan or examination at each collection:
The American Museum of Natural History, New
York, USA (Nancy Simmons); The Carnegie Museum
of Natural History, Pittsburgh, USA (Suzanne
McLaren); The Estación Biológica de Doñana, Se-
villa, Spain (José Cabot); The Croatian Natural His-
tory Museum, Zagreb, Croatia (Igor Pavliniƒ); The
Harrison Institute, Sevenoaks, United Kingdom (Paul
J. J. Bates); Hungarian Museum of Natural History,
Budapest, Hungary (Gábor Csorba); Mammal Re-
search Institute, Polish Academy of Science, Bia»o-
wieóa, Poland (Elwira Szuma); The Museum of
Southwestern Biology, Albuquerque, New Mexico,
USA (William Gannon); The Museum of Texas Tech
University, Lubbock, USA(Robert Baker); The Royal
Ontario Museum, Toronto, Canada (Judith Eger); and
The Zoologisches Forschungsinstitut und Museum
Alexander Koenig, Bonn, Germany (Reiner Hutte-
rer). The senior author was supported by the State
Committee for Scientific Research (Komitet Bada½
Naukowych) – grant no. 6 P04C 07718. The Reflex
Microscope used to measure the fruitbats was kindly
loaned to RDO by Richard Cifelli, University of
Oklahoma, USA. Dr. F. J. Rohlf made a substantial
contribution to this paper by providing detailed com-
ments. In fairness to Rohlf, it should be noted that he
does not agree with the primary conclusions of this
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48 W. Bogdanowicz, J. Juste, R. D. Owen, and A. Sztencel
Specimens used in the present study. Acronyms: AMNH: American Museum of Natural History, New York,
USA; Acronyms: CM — Croatian Natural History Museum, Zagreb, Croatia; CMNH – Carnegie Museum of
Natural History, Pittsburgh, USA; EBD – Estación Biológica de Doñana, Spain; HNHM – Hungarian Museum
of Natural History, Budapest, Hungary; HZM – Harrison Institute, Sevenoaks, United Kingdom; MRI –
Mammal Research Institute, Polish Academy of Science, Bia»owieóa, Poland; MSB – Museum of Southwestern
Biology, Albuquerque, New Mexico, USA; ROM – Royal Ontario Museum, Toronto, Canada; TTU – Texas
Tech University, Lubbock, USA; ZFMK – Zoologisches Forschungsinstitut und Museum Alexander Koenig,
Bonn, Germany. Letters after species number refer to geographic location in the Gulf of Guinea: a – Annobón;
b – Bioko; ca – Cameroon; p – Principe; rm – Río Muni; st – São Tome
Received 18 June 2004, accepted 22 December 2004
Eidolon helvum, XX(n = 44): CMNH40990 – ca;
EBD17386 – st; EBD17387 – st; EBD17388 – st;
EBD17520 – st; EBD17540 – st; EBD17598 – a;
EBD17600 – a; EBD17603 – a; EBD18488 – a;
EBD18825 – b; EBD18826 – b; EBD18827 – b;
EBD18840 – b; EBD18874 – a; EBD18875 – a;
EBD18876 – a; EBD18897 – rm; EBD18898 – rm;
EBD18899 – b; EBD18901 – b; EBD18907 – a;
EBD18913 – st; EBD19044 – b; EBD19064 – b;
EBD19217 – rm; zBD19218 – rm; EBD19224 – rm;
EBD19225 – rm; EBD19226 – rm; EBD19227 – rm;
EBD19238 – rm; EBD20348 – st; EBD20358 – st;
EBD20362 – st; EBD20448 – st; EBD8818 – rm;
EBD8820 – rm; ROM50865 – ca ROM54940 – ca;
ROM55626 – ca; ROM55627 – ca; ZFMK73368 –
Geometric morphometrics and cladistics in mega- and microbats 49
ca; ZFMK74302 – ca; YY (n= 68): EBD17382 – p;
EBD17389 – st; EBD17471 – st; EBD17519 – p;
EBD17538 – p; EBD17539 – p; EBD17542 – st;
EBD17595 – rm; EBD17597 – a; EBD17599 – a;
EBD17601 – a; EBD17602 – a; EBD18199 – p;
EBD18204 – st; EBD18212 – st; EBD18487 – a;
EBD18489 – a; EBD18492 – a; EBD18494 – a;
EBD18828 – b; EBD18829 – b; EBD18833 – b;
EBD18834 – p; EBD18836 – p; EBD18837 – p;
EBD18844 – rm; EBD18869 – p; EBD18879 – b;
EBD18900 – b; EBD18906 – a; EBD18915 – st;
EBD18917 – p; EBD18958 – a; EBD19020 – rm;
EBD19037 – b; EBD19038 – rm; EBD19039 – rm;
EBD19053 – b; EBD19054 – b; EBD19065 – b;
EBD19214 – rm; EBD19215 – rm; EBD19216 – rm;
EBD19220 – rm; EBD19221 – rm; EBD19222 – rm;
EBD19239 – rm; EBD19247 – rm; EBD20344 – st;
EBD20345 – p; EBD20346 – p; EBD20347 – p;
EBD20349 – st; EBD20350 – p; EBD20351 – p;
EBD20352 – p; ; EBD20355 – st; EBD20357 – p; ;
EBD20359 – p; EBD20364 – p; EBD8819 – rm;
EBD8821 – rm; HZM21.4835 – ca; ROM39044 – ca;
ROM39058 – ca; ZFMK64328 – b; ZFMK64331 – b;
ZFMK64333 – b.
Myonycteris brachycephala, XX (n= 9): EBD
18934, 18935, 17410, 17524, 17470, 18904, 17477,
17468, 18936; YY (n= 8): EBD17413, 22284,
17469, 17411, 17526, 18937, 19066, 37413.
Myonycteris torquata, XX (n = 42): AMNH
236237, 236239, 236240, 236246, 236247, 236249,
236254, 236255, 236256, 240999, 241000, 241001,
241002, 241003, 241004; CMNH107996, 40951,
40957; EBD13767, 15046, 15057, 15061, 15065,
15066, 15110, 19058, 22486, 22488, 22489, 22493,
22501, 22503; ROM39393, 43356, 57148, 69001;
TTM17954; TTU17952, 3938, 3939; USNM241112;
ZFMK61621; YY (n = 53): AMNH236236, 236242,
236243, 236245, 236250, 236251, 236252, 236253;
BM139122; CMNH40949, 40950, 40952, 40953,
40954, 40955, 40956, 58253; EBD13888, 15011,
15012, 15013, 15054, 15055, 15056, 15058, 15059,
15060, 15062, 15063, 15727, 17715, 19057, 20487,
20488, 20489, 22487, 22490, 22495, 22497, 22498,
22499, 22504, 22505, 22506, 22507, 22508, 22509,
22510; SNH511901; TTU3937, 3940; ZFMK61623,
Rousettus aegyptiacus, XX (n= 96): AMNH
240988, 240990; CMNH58254; EBD13869, 15152,
15191, 15680, 17352, 17396, 17405, 17406, 17496,
17497, 17533, 17534, 17535, 18203, 18207, 18241,
18250, 18525, 18550, 18557, 18558, 18559, 18841,
18856, 18857, 18858, 18859, 18860, 18861, 18863,
18864, 18865, 18866, 18873, 18918, 18938, 18939,
18952, 19021, 19029, 19030, 19031, 19032, 19033,
19049, 19050, 19052, 19056, 19059, 19267, 19268,
19270, 19276, 19277, 19278, 20178, 20179, 20181,
20183, 20185, 20188, 20190, 20191, 20193, 20196,
20197, 22293, 22307, 22313, 223311; ROM43348,
46756, 55663, 55698, 55699, 55700, 55701, 55702,
55704, 55707, 55732, 55733, 55735, 55737, 56227;
TTM17963; TTU17956, 17962; ZFMK444084,
64312, 64313, 64315, 64319; YY (n = 97): AMNH
240977, 240989, 240996, 318299; CMNH3928; EBD
13679, 15192, 15462, 15463, 17397, 17398, 17521,
17531, 17536, 17597, 17597, 17599, 17601, 17602,
17673, 18208, 18209, 18213, 18242, 18252, 18487,
18489, 18492, 18494, 18830, 18832, 18850, 18851,
18854, 18855, 18865, 18867, 18868, 18870, 18880,
18881, 18891, 18902, 18902, 18906, 18921, 18953,
18954, 18958, 18976, 18978, 18979, 19006, 19007,
19034, 19051, 19060, 19061, 19074, 19269, 19271,
19272, 19273, 19279, 19280, 20180, 20184, 20186,
20195, 20663, 22294, 22308, 22309, 22310, 22312;
HZM504098; ROM46755, 55693, 55694, 55885,
56207, 56226, 56250, 56251, 56255, 58309, 58340;
TTM17958; TTU3923, 3924, 17961; ZFMK64309,
64310, 64314, 64316, 64317, 64318.
Antrozous pallidus, XX (n= 3): ROM67106,
67107, 67111; YY (n= 3): ROM67110, 67121,
Barbastellus barbastellus, XX (n= 3): HNHM
53.29.1, 58.65.1, 2000.43.4; YY (n= 4): HNHM
57.56.1., 57.110.1, 70.18.1; MRI85469.
Barbastella leucomelas, YY (n= 1): HNHM
Corynorhinus mexicanus, YY (n= 2): AMNH
203933, 203934.
Corynorhinus rafinesquii, XX (n= 3): AMNH
142010, 142003, 166892; YY (n= 3): AMNH
142005, 142011, 142006.
Eptesicus fuscus, XX (n= 3): ROM41705,
43176, 46037; YY (n= 3): ROM19388, 24448,
Euderma maculatum, XX (n= 1): MSB96066.
Idionycteris phyllotis, XX (n= 1): AMNH
178893 ; YY (n= 1) AMNH185341.
Myotis lucifugus, XX (n = 3): ROM43605,
88947, 88951; YY (n= 3): ROM40122, 43595,
Otonycteris hemprichi, XX (n= 1): HZM1122.
Plecotus auritus, XX (n= 3): MRI10638, 49960,
57090; YY (n= 3): MRI85094, 91305, 91306.
Plecotus austriacus, XX (n= 3): MRI130/91303,
7868, 96981; YY (n= 3) MRI12410, 38248, 96982.
Plecotus kolombatovici, XX (n= 3): CM3004,
3006, 3008; YY (n= 2): CM2152, 3054.
... Phylogenetic signal was found in almost all the morphogeometric analyses performed here. Similar results were recorded in previous studies focused on other mammalian groups (Bogdanowicz et al., 2005;Cardini and Elton, 2008;Barčiová, 2009). Unlike classical morphometrics, the results obtained here indicate a significant correlation between Procrustes and phylogenetic distances. ...
... Even more, the orthonormal variance decomposition analysis reveals that the cranial morphospaces reflect phylogenetic patterns. Several studies have reported that morphogeometrical cranial data retain phylogenetic information, and it was possible to reconstruct phylogenetic relationships consistent with molecular hypothesis (Polly, 2003;Bogdanowicz et al., 2005;Caumul and Polly, 2005;Cardini and Elton, 2008;Barčiová, 2009;González-José et al., 2008). ...
Three orders of South American extinct native ungulates are recorded from the Santa Cruz Formation along the Atlantic coast of Patagonia: the Notoungulata (Adinotherium, Nesodon, Interatherium, Protypotherium, Hegetotherium and Pachyrukhos), the Litopterna (Theosodon, Anisolophus, Tetramerorhinus, Diadiaphorus and Thoatherium) and the Astrapotheria (Astrapotherium). This work is an ecomorphologic study of these taxa based on geometric morphometrics of the masticatory apparatus. As a reference sample, 618 extant specimens of the orders Artiodactyla, Perissodactyla, Hyracoidea and Diprotodontia were included. Thirty-six cranial and 27 mandibular three-dimensional landmarks were digitized. Allometric scaling, principal component analyses, and phylogenetic generalized estimating equations on the cranium and mandible were preformed separately. The cranial analyses show strong phylogenetic constrains, whereas the mandibular analyses show a functional pattern related to habitat-diet and hypsodonty. The extant brachydont and closed habitat ungulates show a more elongated and narrower mandibular symphysis with a lower mandibular corpus, than hypsodont, open habitat species. The latter have short symphyses with a high, curved mandibular corpus. This general pattern was was also present among Santacrucian ungulates, which permit characterization of the notoungulates mainly as open habitats dwellers, with some foraging on grass (Protypotherium, Interatherium), and others on grass and leaves (Hegetotherium, Pachyrukhos and Adinotherium), depending on the availability. Nesodon may have dwelled in mixed habitats and had a mixed feeding behavior, while small proterotheriids (Anisolophus and Thoatherium) may have fed predominantly on dicotyledonous plants. The remaining litopterns (Tetramerorhinus, Diadiaphorus and Theosodon) and Astrapotherium may have foraged in closed habitats and fed on dicotyledonous plants.
... Geometric morphometrics analysis is a robust tool to highlight interspecific variation and corroborate the phylogenetic relationships within animal groups (e.g. Bogdanowicz et al., 2005;Pavan & Marroig, 2016). In addition, ecological niche studies are being increasingly used for these same purposes (e.g. ...
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A recent phylogenetic analysis of Triatoma pallidipennis, an important Chagas disease vector in Mexico, based on molecular markers, revealed five monophyletic haplogroups with validity as cryptic species. Here we compare T. pallidipennis haplogroups using head and pronotum features, environmental characteristics of their habitats, and ecological niche modeling. To analyze variation in shape, images of the head and pronotum of the specimens were obtained and analyzed using methods based on landmarks and semi-landmarks. Ecological niche models were obtained from occurrence data, as well as a set of bioclimatic variables that characterized the environmental niche of each analyzed haplogroup. Deformation grids for head showed a slight displacement towards posterior region of pre-ocular landmarks. Greatest change in head shape was observed with strong displacement towards anterior region of antenniferous tubercle. Procrustes ANOVA and pairwise comparisons showed differences in mean head shape in almost all haplogroups. However, pairwise comparisons of mean pronotum shape only showed differences among three haplogroups. Correct classification of all haplogroups was not possible using discriminant analysis. Important differences were found among the environmental niches of the analyzed haplogroups. Ecological niche models of each haplogroup did not predict the climatic suitability areas of the other haplogroups, revealing differences in environmental conditions. Significant differences were found between at least two haplogroups, demonstrating distinct environmental preferences among them. Our results show how the analysis of morphometric variation and the characterization of the environmental conditions that define the climatic niche can be used to improve the delimitation of T. pallidipennis haplogroups that constitute cryptic species.
... Geometric morphometric analysis is a robust tool to highlight interspecific variation in zoological groups, such as mammals, corroborating the phylogenetic relationships within them (e. g., Bogdanowicz et al. 2005;Camul and Polly 2005;Pavan and Marroig 2016). In addition, ecological niche studies are being increasingly used for these same purposes (e. g., Rissler and Apodaca 2007; Rivera et al. 2018;Zhao et al. 2019), and for making inferences related to evolutionary questions of both historical distributions and speciation processes (Graham et al. 2004). ...
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The integrative taxonomy approach has recently been widely suggested in systematic studies. Lines of evidence such as the geometric morphometrics and ecological analyses have been useful for discriminating between genetically well-differentiated species. Within the genus Reithrodontomys, R. mexicanus is one of the more taxonomically complex species, being considered a cryptic species complex. R. cherrii was considered a subspecies of R. mexicanus, until molecular evidence raised it to the species-level. Herein, we evaluate these two forms using morphological and ecological data based on the premise that they constitute genetically differentiated species. We carried out geometric morphometric analyses on dorsal and ventral views of the skull. Landmark and semi-landmark configurations for both views of the skull were selected based on previous studies of cricetid rodents. We tested the presence of sexual dimorphism, and the skull shape and size differences between species on both cranial views. Additionally, we characterized the environmental space of each species habitat using bioclimatic variables , elevation, and the Normalized Difference Vegetation Index (NDVI). Females and males of R. mexicanus and R. cherrii did not show sexual dimorphism in shape or size of both skull views. We found significant differences between the two species in both shape and size of the skull. Cranial structures of the ventral view were more useful to differentiate both species. R. mexicanus exhibited a broader environmental space than R. cherrii, with relatively similar values of temperature and elevation, but not of precipitation. The pairwise comparison showed significant differences in the majority of the environmental variables analyzed. Although for each view, we found statistical differences in the skull shape of R. cherrii and R. mexicanus, the ventral side showed major resolutive power differentiating both species. Our findings suggest that R. cherrii tends to have a larger skull than R. mexicanus. However, the morphological and pelage coloration similarity between these species reported in the past, could explain the previous inclusion of R. cherrii as a subspecies of R. mexicanus. R. mexicanus occurs in a variety of vegetation-types coinciding with the broader environmental space that it occupies compared to that of R. cherrii. The natural areas where both species are distributed were associated with high NDVI values. Our results complement the molecular evidence and, under an integrative taxonomy approach, support R. cherrii as a different species from R. mexicanus.
... During the last two decades, several proposals for estimating phylogenies from morphometric data have been discussed contentiously. Some authors have suggested phylogenetic analyses based on cladistic characters derived from partial warp scores Zelditch et al. 1995Zelditch et al. , 1998Swiderski et al. 1998;Bogdanowicz et al. 2005;Clouse et al. 2011) or principal component (PC) scores (MacLeod 2002;González-José et al. 2008Aguilar-Medrano et al. 2011;Brocklehurst et al. 2016). These proposals, however, have been criticized for various reasons, especially the decomposition of phenotypic spaces into distinct characters (Bookstein 1994;Naylor 1996;Adams and Rosenberg 1998;Rohlf 1998;Monteiro 2000;Adams et al. 2011;Zelditch et al. 2012). ...
In recent years, there has been controversy whether multidimensional data such as geometric morphometric data or information on gene expression can be used for estimating phylogenies. This study uses simulations of evolution in multidimensional phenotype spaces to address this question and to identify specific factors that are important for answering it. Most of the simulations use phylogenies with four taxa, so that there are just three possible unrooted trees and the effect of different combinations of branch lengths can be studied systematically. In a comparison of methods, squared-change parsimony performed similarly well as maximum likelihood, and both methods outperformed Wagner and Euclidean parsimony, neighbor-joining and UPGMA. Under an evolutionary model of isotropic Brownian motion, phylogeny can be estimated reliably if dimensionality is high, even with relatively unfavorable combinations of branch lengths. By contrast, if there is phenotypic integration such that most variation is concentrated in one or a few dimensions, the reliability of phylogenetic estimates is severely reduced. Evolutionary models with stabilizing selection also produce highly unreliable estimates, which are little better than picking a phylogenetic tree at random. To examine how these results apply to phylogenies with more than four taxa, we conducted further simulations with up to eight taxa, which indicated that the effects of dimensionality and phenotypic integration extend to more than four taxa, and that convergence among internal nodes may produce additional complications specifically for greater numbers of taxa. Overall, the simulations suggest that multidimensional data, under evolutionary models that are plausible for biological data, do not produce reliable estimates of phylogeny.
... Both PCA and allometric analyses describe a mixture of taxonomic and functional aspects of shape, while the PLS shows an ecomorphological pattern. Although cranial morphology was found more related to taxonomy than ecology (Bogdanowicz et al., 2005;Cardini and Elton, 2008;Barčiová, 2009;Piras et al., 2011;Cassini, 2013;Foth et al., 2015;Murta-Fonseca and Fernandes, 2016), there are many examples of ecomorphological association even during ontogeny (e.g., Merino et al., 2005;Urošević et al., 2013;Segura et al., 2013Segura et al., , 2017Segura, 2015;Olsen, 2017). ...
Ontogenetic variation of cranial characters used in crocodylian phylogenetic systematics has never been studied. Furthermore, the relationship between diet and skull morphological transformation during ontogeny has not been properly explored yet. We quantify the inter- and intraspecific skull morphological variation in extant caiman species focusing on those areas relevant to systematics and, also investigate the relation between diet and morphological changes during ontogeny. We applied a three-dimensional approach of geometric morphometrics on post-hatching ontogenetic cranial series of Caiman latirostris and C. yacare. In order to incorporate incomplete material, we additionally tested four different methods of missing landmark estimation and apply the thin-plate spline interpolation. We detected morphological changes between species and during ontogeny (snout and pterygoid flanges increase their proportions and, orbits, temporal fenestrae, skull roof and foramen magnum decrease their relative size) that constitutes part of a general morphological change in the cranial ontogeny of crocodylians. Moreover, the negative allometry of the fenestrae and neurocranium and the positive allometry of the splanchnocranium in both caiman species are the plesiomorphic condition, at least, for tetrapods. Shape changes during growth were found to be related to ontogenetic changes in the diet. Dissimilarities between species seem to be related to different mechanical requirements and different use of the habitat. We found inter- and intraspecific variation in some morphological characters with systematic implications (the contact of nasals with naris, the contact of prefrontals in the midline, and the bones that border the suborbital fenestra and the proportion in which one of them participates) that are not currently considered in phylogenetic analyses.
... d intra-específica (Dujardin et al. 1997, 1998, Rubio-Pallis 1998, Calle et al. 2002, 2008, Yurtas et al. 2005. Los datos morfométricos también han sido empleados para la postulación de hipótesis cladísticas, demostrando que estos caracteres contribuyen en el apoyo de grupos monofiléticos (Guerrero et al. 2003, Acero et al. 2005, Camul & Polly 2005, Bogdanowicz et. al. 2005, Goloboff et al. 2006, González-José et al. 2008, de Bivort et al. 2010. Más recientemente, Catalano et al. (2010) han desarrollado algoritmos para realizar análisis cladísticos a partir de la inclusión directa sin codificación de las coordenadas x, y (y 3D) de especímenes alineados. ...
Tribe Rhodniini includes Rhodnius Stål and Psammolestes Bergroth. Enzymatic and molecular evidence suggest the tribe is monophyletic. Most species are wild, living in palms and bird nests. Traditionally both genera were considered related; nevertheless, molecular studies don't support the Rhodnius monophyly. The goal was to phylogenetically analyze morphometric variation in wing architecture in support of Rhodniini taxonomy and systematics. We photographed 524 wings of five representatives of Rhodniini: Psammolestes arthuri (Pinto) (n = 89), Rhodnius pictipes Stål (n = 21), R. robustus Larrousse (n = 24), R. prolixus Stål (n = 16), and R. neivai Lent (n = 22). As outgroups we studied four representatives of Triatomini: Eratyrus mucronatus Stål (n = 15), Panstrongylus rufotuberculatus (Champion) (n = 45), P. geniculatus (Latreille) (n = 183), and Triatoma maculata (Erichson) (n = 109). Landmark coordinate (x, y) configurations were registered and aligned by Generalized Procrustes Analysis. Covariance Analyses were implemented with proportions of re-classified groups and MANO VA. Then, wing shape variables (confidence intervals from relative warps) and centroid size were cladistically analysed. Statistical analyses of variance found not significant differences in wing isometric size (Kruskal-Wallis) among P. arthuri-R. neivai-R. pictipes; R. robustus-R. prolixus-T. maculata and between P. rufotuberculatus-P. geniculatus. The a posteriori re-classification was perfect in E. mucrunatus 100% and R. pictipes, followed by T. maculata 96%, R. neivai 95%, P arthuri 93.2%; R. prolixus 87.5%, P. geniculatus 87.4%, P. rufotuberculatus 84.4%, and R. robustus 76%. Cladistic analyses under parsimony selected two most parsimonious trees (L=4.461 IC=0.973 and IR=0.979), where the strict consensus showed a monophyletic group with Panstrongylus (rufotuberculatus + geniculatus) and Triatoma + Rhodniini (Rhodinus + Psammolestes), but internally it shows the paraphyly of Rhodnius regarding Psammolestes. The congruence between these results and previous molecular analyses in Rhodniini, reveal the phylogenetic information of our morphometric characters as support to systematic studies, allowing the combination of geometric morphometrics and phylogenetic methods for the first time in this group.
... Soto-Vivas et al. 2011;Bechara y Liria 2012), así como de vertebrados (e.g. Acero et al. 2005;Cordeiro-Estrela et al. 2008;Marchán-Rivadeneira et al. 2010); en este último caso, se han utilizado en diferentes estructuras morfológicas como cráneos, molares, mandíbulas y escapulas en diversos grupos de mamíferos para determinar variaciones intra e interespecíficas y establecer relaciones filogenéticas (Guerrero et al. 2003;Bogdanowicz et al. 2005;Camul y Polly 2005;Morgan 2009;Lencastre 2011). ...
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In this study, we compared a new population of Rhipidomys from the Sierra de Aroa-Yaracuy State in Venezuela with taxa included within the R. fulviventer section sensu Tribe (1996). We used geometric morphometrics from selected landmarks on the skull, and mandible of three species of Rhipidomys, and three subspecies of R. fulviventer housed in Venezuelan museums. We grouped every taxon as R. venustus Aroa (Sierra de Aroa), R. venustus (Sistema de colinas Lara-Falcón, Cordillera Central, and Andes-Cordillera de Mérida), R. wetzeli (Guayana), R. fulviventer elatturus (Andes-El Tamá), R. fulviventer ssp. 1 (Cordillera Central), and R. fulviventer tenuicauda (Cordillera Oriental). We measured every landmark with its opposite with the purpose to corroborate the results with the linear morphometrics, using the program TPSDig. The results support the proposed taxonomy that previously recognized three species, and three subspecies within R. fulviventer. However, the analysis separated R. venustus Aroa as a distinct group. The landmarks on the dorsal view showed the greatest differences in the Rhipidomys studied. R. venustus Aroa, and R. venustus were the two largest taxa, and R. wetzeli was the smallest. Among the subspecies of R. fulviventer, R. f. ssp. 1, showed less differentiation in the shape, and R. f. tenuicauda showed the strongest differentiation. The Sierra de Aroa population needs careful taxonomic revision because it may represent an undescribed taxon within R. venustus.
... (Rohlf, 2002) . ‫ايهي‬ (Bogdanowicz et al., 2005) . (Rohlf, 2004) . ...
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We applied geometric morphometric techniques to explore the morphological variation of forewings between 10 Asian Calopteryx splendens populations including Azerbaijan, Russia, Turkey, Uzbekistan, Iran, Turkmenistan, Tajikistan, Kazakhstan, and Kyrgyzstan countries. We focused on the study of the phenetic relationships among the populations in central Asia. The results showed that the northern and western populations of Iran had the largest and smallest centroid size of the wings, respectively. In addition, differences among wing shape of the 10 studied populations of C. splendens were significant. Our results indicated that Tajikistan population has quite distinct divergence and also Turkmenistan and northern part of Iran populations both were very close each other and located in a separate clade. The Azerbaijan, Russia, Turkey, Uzbekistan, west Iran, Kazakhstan and Kyrgyzstan populations were revealed to be more interrelated to each other, although Kazakhstan and Kyrgyzstan populations seems to be more closer than the other.
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La Morfometr ía Geométrica es uno de los métodos más avanzados que analizan la forma de estructuras biológicas y ha sido empleada, principalmente, a partir de fotograf ías. Estas representaciones bidimensionales tienen desvent ajas y el uso de modelos 3D supone un avance, que aún no se ha evaluado con precisión. En este trabajo se evalúan los métodos de morfometría bidimensional y tridimensional comparando cráneos en cuatro especies de jutías cubanas (Rodentia, Capromyidae): Mysateles prehensilis , Mesocapromys melanurus, Capromys pilorides y Mesocapromys auritus . Se tomaron fotografías de 64 cráneos en vista lateral, dorsal y ventral, y además se construyeron los modelos 3D de los mismos por métodos de fotogrametría. Se colocaron 54 puntos clave en ambas representaciones de los cráneos para su análisis, y las configuraciones fueron estandarizadas mediante un registro Procrustes. A las variables derivadas se les calcularon los estadísticos descriptivos y se compararon entre las configuraciones 2D y 3D por medio de pruebas de comparaciones pareadas y múltiples de medias empleando métodos de Montecarlo. Además, se compararon las especies mediante diagramas de distorsión para las configuraciones bidimensionales, mientras que para las 3D se realizó mediante mapas de distancias. Se realizaron redes neurales clasificatorias utilizando ambos tipos de datos y se compararon los resultados de las mismas. Los Componentes Principales y los Tamaños de Centroide con las coordenadas 3D mostraron diferencias más claras entre las especies que las variables que utilizaron datos 2D. La Media Procrustes y las distancias entre puntos no diferenciaron claramente entre las especies. Las redes neurales y los análisis de agrupamiento que utilizaron datos tridimensionales fueron más eficaces y eficientes durante la clasificación de especies . Los mapas de distancias 3D mostraron de forma más clara las diferencias entre especies que los diagramas de distorsión. Los modelos tridimensionales presentaron la calidad suficiente como para realizar estudios morfométricos, lo que avala el uso de la fotogrametría en este tipo de trabajos, constituyendo una alternativa más barata que los escáneres 3D. Los resultados mostraron que los análisis de morfometría en 3D resultan mucho más potentes que las 2D, permitiendo diferenciar y caracterizar mejor a cada especie. Geometric Morphometry is one of most advanced methods to analyze shapes of biological structures, and it has been used mainly based on photographs. These bi-dimensional representations have disadvantages so using 3D models should repr esent an advance but it is not fully assessed. In this research, two-dimensional and three-dimensional based morphometric methods are evaluated by comparing skulls in four species of Cuban hutias (Rodentia, Capromyidae): Mysateles prehensilis, Mesocapromys melanurus, Capromys pilorides and Mesocapromys auritus . In 64 skulls we took pictures in dorsal, lat eral and ventral view and 3D models were obtained by photogrammetry techniques. In both representations of the skulls 54 landmarks were placed for their analysis, and the configuration where standardized through Procrustes registration. To the derived variables descriptive statistics were calculated and compared between 2D and 3D configurations by using paired and multiple comparisons tests with Montecarlo methods. The species were compared using distortion diagrams for the two-dimensional configurations, while for 3D models comparison was done using distance maps. Neural classificatory networks were made using both types of data and their results were compared. The Principal Components and the Centroid Sizes of the 3D coordinates found clearer differences bet ween the species than the variables that used 2D data. The variables Average Procrustes and Distances between points did not show clear differences between the species. Neural networks and cluster analyzes that used three-dimensional data were more efficient during the classification of species. The 3D distance maps showed more clearly the differences bet ween species than the distortion diagrams. The threedimensional models obtained show enough quality to perform morphometric studies, which supports the use of photogrammetry in this type of work, been an affordable alternative to expensive 3D scanners. The results showed that 3D morphometric analyses are much more powerful than 2D based, allowing better differentiation and characterization of each species.
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Published studies have shown that methane yield (g CH 4 /kg dry matter) from sheep is positively correlated with the size (volume and surface area) of the reticulo-rumen (RR) and the weight of its contents. However, the relationship between CH 4 yield and RR shape has not been investigated. In this work, shape analysis has been performed on a data set of computerised tomography (CT) scans of the RR from sheep having high and low CH 4 yields ( n =20 and n =17, respectively). The three-dimensional geometries of the RRs were reconstructed from segmented scan data and split into three anatomical regions. An iterative fitting technique combining radial basis functions and principal component (PC) fitting was used to create a set of consistent landmarks which were then used as variables in a PC analysis to identify shape variation within the data. Significant size differences were detected for regions corresponding to the dorsal and ventral compartments between sheep with high and low CH 4 yields. When the analysis was repeated after scaling the geometries to remove the effect of size, there was no significant shape variation correlating with CH 4 yield. The results have demonstrated the feasibility of CT-based computational shape determination for studying the morphological characteristics of the RR and indicate that size, but not shape correlates with CH 4 yield in sheep.
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Phylogenetic relationships among the taxa of plecotine bats (Plecotus, Idionycteris, Barbastella, Euderma, and Corynorhinus) were examined using 32 characters of the skin and skull. Character states for the hypothetical ancestor were inferred by evaluation of outgroup taxa including 12 species of Myotis, two species of Pipistrellus, and Lasionycteris noctivagans. Cladistic analysis performed using PAUP yielded one most-parsimonious tree. The cladogram indicates that each of the taxa is to be regarded as a genus, which supports the contention that Idionycteris is a distinct genus and argues against the subgeneric designation of Corynorhinus. We thus elevate Corynorhinus to full generic status and limit Plecotus to species of the Palearctic.
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Genetic relationships among Rousettus egyptiacus, R. angolensis, Myonycteris torquata, and M. brachycephala were studied based on a starch-gel electrophoretic analysis of 31 presumptive loci encoding 22 enzymatic systems. Data were analyzed by both phenetic and phylogenetic procedures, considering both allele and loci as characters. All of the analyses (both quantitative and qualitative) showed a high level of concordance in establishing R. (Lissonycteris) angolensis, M. torquata, and M. brachycephala as a monophyletic group. Nevertheless, none of the analyses was clearly able to identify sister groups among these taxa. Based on the evidence generated here and from previous independent datasets, the maintenance of Lissonycteris within the genus Rousettus is not sustainable.
The seasonal movements of Eidolon helvum, Myonycteris torquata and Nanonycteris veldkampi were studied. During the dry season, E. helvum roosts in at least one large colony (c500 000 individuals) in the southern forest zone of Ivory Coast. Following the birth of young in February, males and females move into the savanna zones, and the progressive establishment and decline of colonies along a S-N axis indicates that E. helvum migrates at least to the Niger River basin by the middle of the wet season (July). During the dry season, both M. torquata and N. veldkampi are absent from savanna sites, but are common in the forest zone. With the onset of the rainy season in March, catch rates of both species increase first at a southern Guinea savanna site and subsequently at a savanna site 400km to the north, indicating that both species migrate at least to the southern Sudanese savanna zone during this season. Both sexes of N. veldkampi migrate, but the migration of M. torquata is restricted to the immature male cohorts. The high amplitude of the seasonal fluctuations in fruit abundance at savanna sites creates a wet-season surplus of food which results in low intra- and inter-specific competition levels at these sites relative to the forest zone. This may provide the conditions leading to the observed annual migrations.-Author
Limited information from existing data sets and the tremendous amount of diversity in number and kind within the chiropteran family Vespertilionidae (about one-third of all bat species) have hampered efforts to provide adequate assessments of long-standing genealogic hypotheses (e.g., monophyly of the family and of the five subfamilies). We generated approximately 2.6 kilobase pairs of mitochondrial DNA (mtDNA) sequence ecompassing three adjacent genes (12S rRNA, tRNAVa1, 16S rRNA) for 120 vespertilionids representing 110 species, 37 of 44 genera, and all subfamilies. We assessed monophyly of Vespertilionidae in initial analyses of 171 taxa including representatives of all bat families (except the monotypic Craseonycteridae), and assessed lower-level relationships by analysis of several truncated taxon sets. Phylogenetic analysis of ribosomal gene sequences provides well-supported resolution for vespertilionid relationships across taxonomic levels. Furthermore, the resolution is not heavily burdened by alignment of ambiguous regions of the ribosomal gene sequences, and topologies and levels of support produced by two phylogenetic methods (Bayesian and Parsimony) agreed markedly. Our analyses suggest relationships that support many parts of the traditional classification but which also support several changes. The majority of these changes also receives support from other data sources, particularly bacular and karyotypic data. We make more than 20 taxonomic conclusions or recommendations and construct a working classification for vespertilionoid bats. Highlights include: Miniopterus (subfamily Miniopterinae) is recognized in its own family, Miniopteridae, as it represents an extremely divergent lineage relative to other vespertilionids, and in some analyses is sister to the molossids and natalids; all other vespertilionids examined form a well-supported clade; two of the traditional subfamilies within Vespertilionidae (sensu stricto) are monophyletic, Murininae and Kerivoulinae; Nyctophilinac has no validity and Vespertilioninae is paraphyletic relative to the position of Myotis; Myotis is sister to a clade containing Kerivoulinae and Murininae and is recognized in its own subfamily, Myotinae; Myotis subgenera Leuconoe, Selysius, and Myotis are polyphyletic, and a subgeneric classification reflecting geography is suggested, broadening subgenus Myotis to include the sampled Old World species, and allocating the sampled New World species to another subgenus (Aeorestes Fitzinger, 1870); Vespertilioninae (excluding Myotis) is monophyletic; Pipistrellus-like bats (i.e., the traditional tribe Vespertilionini) are divided into three tribes (Nycticeiini, Pipistrellini; Vespertilionini); and support for three tribes of Pipistrellus-like bats has several implications at the genus level. Overall, this study offers a robust working hypothesis for vespertilionid relationships and provides a good starting point for new investigations into the evolutionary history of Vespertilionidae.
It has been claimed that quantified features are inappropriate for phylogenetic analysis. We consider that claim to be true under most conditions for characters discovered by commonly used morphometric methods, including outline-based and conventional multivariate methods. The most important reason these characters are unsuitable is that one of the tests of homology, the test of similarity, may be difficult to apply to them. This test is not even possible if the methods for comparing forms, such as outline-based techniques, do not ensure that the characters are located in the same part of the anatomy. Conventional methods, including principal components analysis, have no explicit basis for localizing characters. In addition, unless the transformation between forms is homogeneous, conventional methods cannot dissect transformations region by region to discover characters. However, one morphometric method, the thin-plate spline decomposed by its partial warps (TPS), finds characters that can be subjected to the same tests of homology (conjunction, similarity, and congruence) that we would apply to all other characters. Among available methods, TPS is unique in being able to locate the center and spatial extent of regional differences in shape and ensures that the same regions are compared among forms. We provide an example using the teleost fishes piranhas, in which tests of homology are applied to a synapomorphy found by the method.