Environmental Skeptics and Critics

Vol. 3, No. 1, 1 March 2014

International Academy of Ecology and Environmental Sciences

Environmental Skeptics and Critics
ISSN 2224-4263
Volume 3, Number 1, 1 March 2014

Editor-in-Chief
WenJun Zhang
Sun Yat-sen University, China
International Academy of Ecology and Environmental Sciences, Hong Kong
E-mail: zhwj@mail.sysu.edu.cn, wjzhang@iaees.org

Editorial Board
Taicheng An (Guangzhou Institute of Geochemistry, Chinese Academy of Sciences,
China)
Andre Bianconi (Sao Paulo State University (Unesp), Brazil)
Alessandro Ferrarini (University of Parma, Italy)
Gianluigi de Gennaro (University of Bari 'A. Moro', Italy)
Marcello Iriti (Milan State University, Italy)
Suyash Kumar (Govt. PG Science College, India)
GuangHua Liu (Guangdong AIB Polytech College, China)
T.N. Manohara (Rain Forest Research Institute, India)
Lev V. Nedorezov (University of Nova Gorica, Slovenia)
Edoardo Puglisi (Università Cattolica del Sacro Cuore, Italy)
Mohammad Hossein Sayadi Anari (University of Birjand, Iran)
Mohammed Rafi G. Sayyed (Poona College, India)
R.N. Tiwari (Govt. P.G.Science College, India)

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Environmental Skeptics and Critics, 2014, 3(1): 1-7

Article

First evidences of sexual divergences in flight behaviour and space

use of lesser kestrel Falco naumanni

Marco Gustin1, Alessandro Ferrarini2, Giuseppe Giglio1, Stefania Caterina Pellegrino1, Annagrazia
Frassanito3
1
Lega Italiana Protezione Uccelli, Conservation Department, Via Udine 3, I-43100 Parma, Italy
2
Department of Evolutionary and Functional Biology, University of Parma, Via G. Saragat 4, I- 43100 Parma, Italy
3
Alta Murgia National Park, via Firenze 10, 70024, Gravina in Puglia, Bari, Italy
E-mail: sgtpm@libero.it,alessandro.ferrarini@unipr.it

Received 30 November 2013; Accepted 5 January 2014; Published online 1 March 2014

Abstract
We present here the first description of recorded sexual differences in flight behaviour and space use of lesser
kestrel Falco naumanni. Lesser kestrel is a migratory, colonial, small falcon breeding mainly in holes and
crevices in large historic buildings within towns and villages, or in abandoned farm houses across the
countryside. Using accurate GPS data-loggers, we gathered data on the activities of lesser kestrels in the two of
main colonies of lesser kestrels in Italy, i.e. Gravina in Puglia and Altamura (Apulia, Southern Italy) and the
surrounding rural areas in a 20-days monitoring during the reproductive period. We tested for sex differences
in space use (home range’s circularity ratio) and flight attributes (5-minute flight length, instantaneous speed,
distance from nest, flight altitude above ground level) of 9 monitored individuals (4 males and 5 females). We
found significant sexual differences for all the observed traits. Our results demonstrate that female lesser
kestrels during the monitoring period employed a lower amount of energy in local movements as measured by
four flight attributes that resulted significantly different (and lower) than for males. Compact home ranges for
females could represent a maximization of the benefit-cost ratio between prospected surface and distance from
nest, i.e. the optimal trade-off between foraging requirements (explored surface) and costs in terms of time and
energy (distance from nest). On the contrary, males showed a significantly different space use with very
elongated home ranges and mean distance from nest almost three times as elevated as females’ one. We argue
that the detected sexual divergence was the product of their respective ways to optimize the relationship
between resource acquisition and reproductive activity.

Keywords data-loggers; flight attributes; local-scale movements; reproductive period.

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2 Environmental Skeptics and Critics, 2014, 3(1): 1-7

1 Introduction
Lesser kestrel Falco naumanni is a migratory, colonial, small falcon breeding mainly in holes and crevices in
large historic buildings within towns and villages, or in abandoned farm houses in the countryside (Negro,
1997). Today the species is considered “least concern” in western Europe (BirdLife International, 2013).
In birds, the sexes usually differ in size if not also in proportions of body parts, being this usually
considered as a consequence of variation among species in social mating system and the pattern of parental
care (Andersson, 1994). However, there is only an occasional reference in the literature to sexual differences in
relation to flight behaviour and space use (Selander, 1966; Weimerskirch et al., 2000), and to date none has
been stated about Falco naumanni.
For this reason, the aim of our work has been to acquire knowledge about sexual differences in flight
behaviour and space use of lesser kestrels during the reproductive period. In order to do this, we: 1) gathered
data on the activities of male and female lesser kestrels; 2) compared the activity of male and female lesser
kestrels with regard to space use and flight attributes. We investigated birds from the two main colonies of
lesser kestrels in Italy, i.e. Gravina in Puglia and Altamura (Apulia, Southern Italy) and the surrounding rural
areas (Bux et al., 2008).

2 Methods
The study area corresponds to the National Park Alta Murgia and the SPA (Special Protection Area) “Murgia
Alta” IT9120007 (Apulia, Southern Italy) and is included within the IBA (Important Bird Area) “Murge”
(Heath and Evans, 2000). It comprehends one of the main colonies of lesser kestrels in Italy (Bux et al., 2008),
i.e. Gravina in Puglia, Altamura and the surrounding rural areas.
Surveys were conducted using TechnoSmart GiPSy-4 data-loggers (23×15×6 mm, 1.8 g plus 3.2 g
battery), that provided information about date, time, latitude, longitude, altitude (meters a.s.l.) and
instantaneous speed (km/h).
Nine individuals (4 males and 5 females) were surveyed for 20 consecutive days from June 21th to July 9th
2012, i.e. the period when youngs were in nests. Eight out of 9 individuals were from Gravina in Puglia, the
other one from Altamura. Data acquisition occurred every 5 minutes during two time periods: day (08:00-
19:00 H) and night (02:00-06:00 H).
GPS data were imported into the GIS GRASS (Neteler and Mitasova, 2008) and added to further layers: a)
boundaries of the Alta Murgia National Park, b) digital terrain model of the study area, c) nest and roost
locations. The digital terrain model at 1:10,000 scale of the study area was digitized by the authors from
available topographic maps of Apulia Region (Fig. 1).
For each GPS point, flight height above ground level was achieved by subtracting terrain elevation from
altitude a.s.l. provided by data-loggers. Five-minute flight length was calculated as length of flight between 2
successive GPS acquisitions. For each GPS point of each individual we also calculated the distance in meters
from its nest.
We estimated home-range (HR) for each individual using the minimum convex polygon method (Graham,
1972). Besides the 100% contour, we also considered the 99% isopleth using probability polygons with the
“proximity to mean of all fixes” criterion (Kenward, 1987).
Last, we used the circularity ratio index (CR; Jennrich and Turner, 1969) as indicator of lesser kestrels’
space use. For each one of the above mentioned HRs, we calculated CR as follows:

S HR
CR  (1)
Scc

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 Environmental Skeptics and Critics, 2014, 3(1): 1-7 3

where SHR is the HR size (in hectares), while Scc is the size (in hectares) of the smallest circumscribing circle.
CR is a unitless measure approximating 1 for compact HRs (similar to circles), and 0 for very elongated ones.
For the purpose of our work, CR was more suitable than HR size. In fact higher values of CR suggest that
individuals prospected larger surfaces whilst limiting their distance from nest, being the opposite for lower
values.

Fig. 1 The digital terrain model at 1:10,000 scale of the study area digitized by the authors from available topographic maps of
Apulia Region. Black and grey points show the presence of nests and roosts in the two colonies of Gravina in Puglia (on the left)
and Altamura (on the right).

Based on descriptive statistics of flight attributes (5-minute flight length, instantaneous speed,
distance from nest, flight altitude above ground level) and space use (CR), we compared males and females
using inferential tests. After testing for normality (Shapiro-Wilk test) and homoscedasticity (Levene's test), we
applied the correct inferential test (T-test or Mann-Whitney test). All the statistical analyses were performed
using SPSS (SPSS Inc., 2007) software and considered significant for p<0.05.

3 Results
The monitoring period amounted to 311 hours, of which 116 hours (1389 GPS points) for females and 195
(2337 GPS points) for males. Lesser kestrels flew 3674.2 km in total, of which 966.5 due to females and
2707.7 to males (Fig. 2).
Descriptive statistics (Tab. 1) suggested differences in flight attributes, hence we tested if:
a) 5-minute flight length for males was significantly higher than for females;
b) instantaneous speed for males was significantly higher than for females;

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4 Environmental Skeptics and Critics, 2014, 3(1): 1-7

c) distance from nest for males was significantly higher than for females;
d) flight altitude above ground level for males was significantly higher than for females.

Fig. 2 Study area (Gravina in Puglia on the left, Altamura on the right and Alta Murgia National Park; Italy) with male and
female flight trajectories followed by 9 individuals of lesser kestrels surveyed from June 21th to July 10th 2012.

We achieved the following results (Table 2): a) positive (p<0.001), b) positive (p<0.001), c) positive
(p<0.001), d) positive (p<0.001). CRs of detected HRs (Fig. 3) resulted significantly higher for females
(p<0.05) for all the considered isopleths (Table 3).

Table 1 Descriptive statistics of flight attributes of female (N1: 1389 GPS points) and male (N2: 2337 GPS points) lesser kestrels.

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 Environmental Skeptics and Critics, 2014, 3(1): 1-7 5

Table 2 Inferential statistics for male and female flight attributes of surveyed lesser kestrels.

Fig. 3 Home-ranges (100% isopleths) of 4 male and 5 female lesser kestrels surveyed from June 21th to July 10th 2012.

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6 Environmental Skeptics and Critics, 2014, 3(1): 1-7

Table 3 Inferential statistics for circularity ratios of female and male lesser kestrels’ home-ranges (100% and 99% isopleths).

4 Discussion and Conclusions

Our work is the first description of recorded sexual divergences of lesser kestrels with regard to local-scale
movements and space use.
We might expect that any sex divergence in lesser kestrel’s behaviour was the product of their respective
ways to optimize the relationship between resource acquisition and reproductive activity. In most animal
species, while egg production is energetically expensive, the gametic contribution of the male usually requires
less energy (e.g., Emlen and Oring, 1977), hence strong sexual divergences in reproductive role were expected
to translate into strong divergences in movements patterns and space use between males and females, at least
during the reproductive season (during 2012, surveyed nests of 77 lesser kestrels in the study area had a clutch
of 3.79 ± 0.82 eggs; Gustin et al., 2012).
Our results demonstrate that female lesser kestrels during the monitoring period employed a lower amount
of energy in local movements as measured by four flight attributes that resulted significantly different (and
lower) than for males. Furthermore, female lesser kestrels exhibited significantly higher CRs. We argue it was
as a consequence of two necessities: a) spending as much time as possible in parental care, b) limiting energy

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 Environmental Skeptics and Critics, 2014, 3(1): 1-7 7

requirements for resource acquisition. Owing to budgetary requirements of time and energy for reproduction
and parental care, an upper limit to female flight activities was expected.
Compact HRs for females represented a maximization of the benefit-cost ratio between prospected surface
and distance from nest, i.e. the optimal trade-off between foraging requirements (explored surface) and costs in
terms of time and energy (distance from nest). On the contrary, males showed a significantly different space
use with very elongated HRs and mean distance from nest almost three times as elevated as females’ one.
Possible reasons for these long-distance flights include their roles as an exploratory surveillance of the HR, the
search for food-rich patches and exploring neighbouring territories (Perez-Garcia et al., 2012).
In order to detect if sexual differences in flight behaviour and space use of lesser kestrels is a prerogative
of the reproductive period, we are planning to extend our surveys to the pre-reproductive period.

Acknowledgements
We thank the Alta Murgia National Park that funded this study (grant number 2011/381). We thank Dr. Ugo
Mellone (University of Alicante) and Dr. Mattia Brambilla (Fondazione Lombardia per l’Ambiente) for their
useful suggestions.

References
Andersson M. 1994. Sexual Selection. Princeton University Press, Princeton, USA
BirdLife International. 2013. Species factsheet: Falco naumanni. http://www.birdlife.org
Bux M, Giglio G, Gustin M. 2008. Nest box provision for lesser kestrel Falco naumanni populations in the
Apulia region of southern Italy. Conservation Evidence 5: 58-61
Emlen S, Oring L. 1977. Ecology, sexual selection and the evolution of mating systems. Science, 197: 215-223
Graham R. 1972. An efficient algorithm for determining the convex hull of a finite point set. Information
Processing Letters 1: 132-133
Gustin M, Ferrarini A, Giglio P, Pellegrino S, et al. 2012. Il Parco per il grillaio (Falco naumanni) nel Parco
Nazionale dell’Alta Murgia. Recupero pulli, divulgazione e monitoraggio. Technical Relation (in Italian)
Heath MF, Evans M.I. 2000. Important Bird Areas in Europe: Priority Sites for Conservation. BirdLife
International, Cambridge, UK
Jennrich RI, Turner FB. 1969. Measurement of non-circular home range. Journal of Theoretical Biology, 22:
227-237
Kenward R. 1987. Wildlife Radio Tagging. Academic Press, London, UK
Negro JJ. 1997. Falco naumanni Lesser Kestrel. Birds of the Western Palearctic Update, 1: 49-56
Neteler M, Mitasova H. 2008. Open Source GIS: A GRASS GIS Approach. Springer, New York, USA
Perez-García JM, Margalida A, Afonso I, Ferreiro E, et al. 2013. Interannual home-range variation,
territoriality and overlap in breeding Bonelli’s Eagles (Aquila fasciata) tracked by GPS satellite telemetry.
Journal of Ornithology, 154: 63-71
Selander RK. 1966. Sexual dimorphism and differential niche utilization in birds. The Condor, 2: 113-151
SPSS Inc. Released 2007. SPSS for Windows, Version 16.0. SPSS Inc., Chicago, USA
Weimerskirch H, Cherel Y, CuenotChaillet F, Ridoux V. 1997. Alternative foraging strategies and resource
allocation by male and female wandering albatrosses. Ecology, 78: 2051-2063

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Environmental Skeptics and Critics, 2014, 3(1): 8-16

Article

Invasions, adaptive radiations, and the generation of biodiversity

John C. Briggs
Department of Fisheries and Wildlife, Oregon State University, Corvallis, OR 97333, USA. Present address: 2320 Guerneville
Rd., Santa Rosa, CA 95403, USA
E-mail: clingfishes@yahoo.com

Received 3 November 2013; Accepted 8 December 2013; Published online 1 March 2014

Abstract
When the subject of global biodiversity and its loss through human action became a focal point of conservation
biology, there developed an increasing argument about the effects of the invasions of exotic species into native
ecosystems. In order to place the current contention in a historical context, evidence from the geologic record
has been examined. The record clearly indicates that numerous species invasions, extending from the
Paleozoic through the Cenozoic, have led to adaptive radiations significantly increasing global biodiversity.
Three invasion induced categories are recognized: (1) global radiations, (2) local radiations, and (3) single
species effects. Together, the three may account for much of the biodiversity gain that has accumulated in the
intervals between mass extinctions. In contemporary time, it has become apparent that exotic species
colonizing a native ecosystem rarely cause extinctions. Instead, the invaders are accommodated by the native
species that occupy the appropriate niches or habitats. The accommodation process results in a gain in the
species diversity of the invaded area. Over time, local diversity gains can result in global gains as speciation
among the invaders takes place. The majority of successful invasions occur via migration of species from
centers of high species diversity to places of lesser diversity. The result is a dynamic world characterized by
constant movement whereby the high diversity centers increase the diversity of outlying regions.

Keywords adaptive radiation; biodiversity; evolution; historic invasions; species invasions.

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1 Introduction
Adaptive radiation may be defined as the diversification of species to fill a wide variety of ecological niches. It
occurs when a single ancestral species gives rise, through repeated episodes of speciation, to numerous kinds
of descendents that remain or become sympatric (Lomolino et al., 2010). Although the initial cause is often
attributed to development of a key innovation in the ancestral species, that appears to be a simple solution to a
more complex process. Simpson (1953) proposed entry into an adaptive zone as a prerequisite for adaptive
radiation. An adaptive zone could be entered in one of three ways: (1) evolution of a key innovation, (2)

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 Environmental Skeptics and Critics, 2014, 3(1): 8-16 9

dispersal into a new habitat, and (3) extinction of antagonists. More recently, the causes and significance of the
adaptive radiation phenomenon have been widely discussed. Prominent contributions have been a book by
Schluter (2000) and a detailed review by Yoder et al. (2010). Currently, such radiations are most commonly
attributed to ecological opportunity, meaning a relaxation of selection or an ecological release. While these
general terms cover many possibilities, I find Simpson’s original subdivisions still appear to be logical and
useful.
An important question is, which of the three main causes of adaptive radiation (key innovation, invasion
into a new habitat, extinction of competition) is the most important driver of biodiversity increase? This
question has not been asked previously because the relationship between adaptive radiations and global
biodiversity had not been explored in detail. Judging from observations in the paleontological literature, as
well as those on contemporary species, it appears that species invasions are the initial step that precedes
evolutionary change. Once a given species finds an advantageous new habitat, it begins the adaptation process
which may result in significant or key innovations. Instead of one important driver, there is really a sequence:
invasion to evolutionary innovation to radiation. The rapidity and the ultimate extent of the radiation may be
influenced by the residual effects of previous extinctions.
Although the term “biodiversity” generally means the diversity of life in the broad sense, it is here
employed as a synonym for species richness or species diversity. This is consistent with the meaning of the
term in most works dealing with conservation issues. Species that colonize a new ecosystem or environment,
as the result of natural or human introductions, are characterized as invasive. Otherwise, they possess no
special attributes and may be undesirable or beneficial according to various economic or esthetic standards.
Each species needs to be judged on its own merits, not whether it happens to be exotic or native (Davis et al.,
2011). Invasive species, whether entering a new unoccupied environment or one that is well populated, appear
to follow the sequence noted above. If the environment is unoccupied, there would no native opposition and
the invasion would proceed rapidly.
The prevention of biodiversity loss has become the foremost conservation goal of the 21st century. It is the
primary concern of many scientists and the focus of action among most conservation societies. The purpose of
this paper is to first, explore the biodiversity effects of historic invasions and second, to compare those results
to recent knowledge gained about contemporary invasions. The comparison process provides evidence bearing
on the vital question of loss or gain in global biodiversity.

2 Historic Global Invasions

Although the following invasions have all been identified in the fossil record, new information exists about
most of them and listing them, in order of habitat invaded, should provide a useful comparison when
evaluating Recent events. That is, current invasions may take on an added importance when viewed through
the lens of history. Information is presented in a sequence: organisms - origin - invasion - time - reference -
remarks.
1. Green algae - freshwater - land - 468 to 472 Ma (million years ago) - Rubenstein et al., 2010.
Perhaps the most important invasion in Earth history. Locality was in Gondwana (Argentina).
2. Progymnosperm seed ferns - swamp - dry land - 385 Ma - Taylor and Taylor, 1993. Gave rise to sister
clades of gymnosperms and angiosperms.
3. Angiosperms - equatorial region - high latitudes - 140 Ma - Magallon and Castillo, 2009; Benton,
2010. Major diversity increase took place 125 - 80 Ma.
4. Insects (Hexapoda) - sea - land - 433 Ma - Wheat and Whalberg, 2013. Rapid radiation to possibly
two million species.

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5. Tetrapod amphibians - aquatic - land - 395 Ma - Niedzwiedzki et al., 2010; Clack, 2012. Clack’s
review notes the many morphological changes that lobe-fin fishes underwent in order to become adapted to
terrestrial life. There is no indication of a key innovation that provided the ability to climb out of the water and
survive on land. Brief subaerial exposures (invasions) were apparently followed by a multitude of innovations
that provided an increasingly better survival on land. As indicated in the introduction, the process consisted of
a sequence: invasion to innovation to radiation.
6. Amniotes - swamp - dry land - 318 Ma - Laurin and Gauthier, 1996. The early amniotes diverged into
two lines, one line (Synapsida) culminated in living mammals, and another line (Sauropsida) embraced all
living reptiles (including birds). The evolution of the amniotic egg was the ecological equivalent of the
development of the seed in vascular plants. In each case, protection of the early embryo from excessive water
loss permitted a great invasion into dry, upland habitats.
7. Insects - land - air - 325 Ma - Wheat and Whalberg, 2013. The adoption of flight, and the numerous
morphological and physiological changes it required, marked the next invasion into a foreign habitat. It was
successful to the extent that more than 98% of modern insects have wings.
8. Pterosaurs - land - air - 251 Ma - Hone, 2012. Pterosaurs, the largest flying animals of all time, were
extant from the Triassic until the Cretaceous (251 – 66 Ma). Their decline has been attributed to the rise of the
birds, but there seems to be no evidence of direct competition (Prentice et al., 2011).
9. Birds - land - air - late Jurassic? - Brocklehurst et al., 2012; Jetz et al., 2012. There was a strong
increase in diversification rate from about 50 Ma to the present.
10. Bats - land - air - 52 Ma - Simmons et al., 2008. Bats (Chiroptera) represent one of the largest and
most diverse orders of mammals, accounting for about one-fifth of extant species.
11. Monocotyledon plants - freshwater - sea - 70 Ma - Vermeij, 2004. About 60 extant species.
12. Marine reptiles - land - sea - 250 Ma - Thorne et al., 2011. A major radiation resulted in the long-
necked fish-eating eosauropterygians, mollusk-eating placodonts, serpentine thalattosaurs, and stream-lined
ichthyosaurs. Marine crocodilians appeared in the Jurassic and mosasaurs in the late Cretaceous (85 Ma).
Marine turtles appeared about 150 Ma (Bowen et al., 1993).
13. Sea snakes (Elapidae, Hydrophiinae) - land - sea - 10 Ma - Sanders et al., 2008. The subfamily
includes more than 100 terrestrial species plus 60 species of completely aquatic sea snakes. The entire
radiation took place within the past 10 Ma. Even more remarkable, the genus Hydrophis, with more than 40
species, evolved within the past 5 Ma.
14. Marine mammals - land - sea - 55Ma - Uhen, 2007; Berta, 2012. Mammals did not enter the sea until
they had evolved on land for about 300 Myr (million years). Four different lineages (Cetacea, Sirenia,
Desmostylia, Pinnipedia) invaded the sea in the Lower Eocene. The whales (Cetacea) evolved into about 70
genera within the following 50 Myr. Living whales include 83 species, pinnipeds 33 species, sirenians 4, and
desmostylians 0.
15. Marine insects - freshwater - sea - 45 Ma - Andersen and Cheng, 2004. The genus Halobates includes
five species of sea striders that are completely oceanic.

3 Global Consequences
Although species diversity in the marine environment showed considerable gains from the Mesozoic through
the Cenozoic, despite the loss at the K/T boundary, it failed to keep up with the enormous increase in terrestrial
diversity that began in the mid-Cretaceous period. The Cretaceous Terrestrial Revolution (CTR) (Benton, 2010)
took place as the angiosperm flora invaded the higher latitudes about 125-80 Ma (Magallo and Castillo, 2009).
As the angiosperms dispersed, the advanced eudicots underwent an explosive radiation (Heimhofer et al., 2005)

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 Environmental Skeptics and Critics, 2014, 3(1): 8-16 11

causing a huge increase in primary productivity (Brodripp and Feild, 2010). Beginning about 100 Ma, the
eudicot clade increased its biomass production three to four times over that of their earlier relatives (Boyce et
al., 2009).
The expansion of angiosperms was accompanied by huge radiations in insects, especially among the
genera of Coleoptera, Hymenoptera, and Lepidoptera, along with comparable increases in plant-associated
fungi (Vermeij and Grosberg, 2010). The productivity increase of the CTR also had notable effects on
vertebrate radiation. For example, the four principal mammalian groups (Marsupialia, Laurasiatheria,
Euarchontoglires, Afrotheria) diverged around 90-80 Ma (Meredith et al., 2011). It was during the CTR that
biodiversity on land finally exceeded that in the sea, an event called “The Great Divergence” (Vermeij and
Grosberg, 2010). Today, there are far more species living on land; in the sea, non-microbial species account
for a small fraction of the global total, estimated at anywhere from 2% (Briggs, 1994) to 5% (Benton, 2009) to
15% (May, 1994).
Mass extinctions appear to cause a collapse of ecospace which must be rebuilt to create new opportunities
and redirect the course of evolution. After an extinction, there is usually a survival interval during which there
is a lack of speciation. This interval is generally followed by a rapid innovation, often consisting of “bloom
taxa” that abruptly diversify and then decline. After the K/T extinction, angiosperm plants began to recover
their diversity after about 1.5 Myr, and marine productivity returned after a few hundred thousand years
(Erwin, 2001). In mammals, several crown-group orders, including Primates and Rodentia, evolved before the
K/T boundary, some rose in the Eocene- such as the Perissodactyla and Carnivora- and others in the later
Cenozoic. But these diversifications were spread out over considerable time and were not as dramatic as those
of the CTR (Meredith et al., 2011). In regard to global biodiversity, it may be observed that mass extinctions
produce losses that are not recovered for hundreds of thousands or millions of years. But in the eventual
aftermath, rapid speciation takes place in some regions that may effect others by means of species invasion, as
will be noted in the case of Cenozoic mammals.
There has been considerable speculation about the evolutionary effects of mass extinctions. These once
provided a consistent message that a “wiping out of the old forms to make way for the new” had evolutionary
benefits. Eldredge (1987) expressed the firm belief that, without extinctions to free up ecological niches, life
would still be confined to a primitive state somewhere on the sea bottom. Stanley (1987) stated, “Had the
dinosaurs survived, there is no question that we would not walk the earth today. Mammals would still be small
and unobtrusive, not unlike the rodents of the modern world.” The idea that major extinctions convey
evolutionary benefits was developed into a new theory of evolution by Hsü (1986). He would substitute the
concept of evolution by means of global extinctions for Darwin’s mechanism of natural selection. In his “new
catastrophism” evolutionary advances would take place as survivors adapted to spaces created by extinction
events. In his view, “it is time to wake up to the absurdity of the idea of natural selection.” Recent advances in
paleontology have made these ideas of the 1980s untenable.

4 Historic Regional Invasions

Some palaeontological works have dealt with regional invasions and may have relevance to contemporary
events. Patzkowsky and Holland (2007) investigated a Late Ordovician marine biotic invasion linked to a
warming event. They found the increase in species diversity caused by the invasion was not ephemeral but
lasted for at least 1 Myr. Another Ordovician study (Heim, 2008) found evidence of a global richness
equilibrium dictated by a combination of invasion, origination, and extinction rates. But, Devonian studies
(Stigall and Liberman, 2006; Stigall, 2012) found a long-term biodiversity decline combined with a
preferential survival of invaders and a drop in speciation rate.

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In regard to the fossil evidence, it seems to me that recent (Cenozoic) data are more applicable to current
invasions than events that took place in the Paleozoic some 300 to 500 Ma. There is the example of the trans-
Arctic invasion of molluscs between the North Pacific and the North Atlantic (Vermeij, 2004). That migration
took place in the Pliocene about 3.5 Ma and the results are apparent in the living fauna. Invasion into the
Atlantic involved 265 species and 24 species invaded the North Pacific. The molluscan fauna of the North
Atlantic now consists of large numbers of species that have evolved from Pacific ancestors. In the
Northwestern Atlantic,118 species or 45% of the molluscs have such ancestry, indicating a long-term effect of
invasions and, via speciation, an increase of global biodiversity. The superior diversity of the Pacific (twice as
many molluscan species) and its greater resistance to invasion probably accounted for the difference in
invasion success. In each recipient area there were no native extinctions attributable to the invaders.
Another case involves the Miocene to early Pleiostocene molluscan invasions between the Caribbean and
Florida (Vermeij, 2005). Some 40 species invaded from the Caribbean, which possessed the richest molluscan
fauna, but only four moved in the opposite direction. During the Pliocene, the invasion accounted for almost
all of the increase in standing diversity in Florida, and about 90% of the invaders underwent speciation. The
long-term consequences of this and many other regional exchanges were enrichment of the local and global
species pool.
Aside from the marine examples, there is an extensive literature devoted regional invasions and adaptive
radiations by birds, mammals, and amphibians. In regard to mammals, the details of their Cenozoic increase in
diversity indicate more than a simple replacement for the dinosaurs. Mammals tended make evolutionary
advances in certain regions and then invaded others where they underwent rapid diversification. For example,
placental mammals invaded South America from North America, marsupials and rodents invaded Australia
from South America and Asia respectively, various African mammals invaded Eurasia, and North American
ungulates reached Eurasia (Lomolino et al., 2010). Similar examples may be found among the amphibians
where the advanced frogs (neobatrachians) may have originated in Africa-India, salamanders in East Asia, and
the caecilians in Triassic Pangaea (Zhang et al., 2005). In regard to birds, rapid local radiations took place in
Asia, North America, and southern South America (Jetz et al., 2012). All of these regional invasions and
numerous others resulted in immediate gains in local biodiversity, followed by speciation providing long-term
gains in global biodiversity.

5 Contemporary Invasions
A review of contemporary invasion ecology was recently published in this journal (Briggs, 2013). It was
concluded that, on land and in the sea, invader species added to local diversity. In the sea, it seemed that the
addition process must have a competitive component because successful (colonizing) invasions took place
almost entirely from areas of high species diversity to those of lesser diversity. On land, various kinds of plant
competition had been described but the relationship between regional and local species diversity was not quite
as clear. But almost all successful invasions were apparently made possible by accommodations on the part of
the native species. The accommodation process can include the provision of space, shelter, or physical support
(Briggs, 2010). Although some reference was made to fossil evidence, space did not permit a detailed analysis.

6 Current Opinions
Although the geologic record indicates that biological invasions are a driving force for global biodiversity,
such a conclusion may seem incongruous in the light of many current opinions on species invasions. During
the past 20 years, invasive species have usually been described in detrimental terms. Queries to yahoo or
google about invasive species will bring up a multitude of postings by a variety of organizations, including the

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 Environmental Skeptics and Critics, 2014, 3(1): 8-16 13

United Nations (IUCN), Wikipedia, Encyclopedia of Earth, UN Environment Program, Global Issues, The
World Bank, Institute for European Environmental Policy, U.S. Environmental Protection Agency, Foundation
for Research and Sustainable Development, Eco-Question, Green Facts, Eco-Pros, and Actionbioscience. In
addition, many individual nations and conservation organizations have web sites emphasizing the danger of
invasive species. These sources provide a consistent message saying that nonnative species cause native
extinctions resulting in biodiversity loss, as well as various kinds of ecological damage. The organizations and
individuals that continue to bemoan biodiversity loss are misleading the public and are directing conservation
support away from the real problem, the precarious existence of small populations that are the results of human
habitat destruction and over exploitation.

7 Discussion
In light of the present day concern about invasion-caused extinctions and loss of biodiversity, it seemed
worthwhile to examine evidence about historic invasions and their biodiversity effects. The greatest adaptive
radiations were caused by several invasions that took place in the early Paleozoic. Most of the terrestrial
world’s present biodiversity can be traced back to the invasions of green plants about 470 Ma, the insects
about 433 Ma, and the amphibians about 395 Ma. Beyond these beginnings, there have been numerous
subsequent invasions and radiations that have also involved large portions of the globe and, through the
millennia, have continued to add to the general biodiversity increase.
After the early Paleozoic, two of the most notable events were invasions of the dry parts of the Earth made
possible for: (1) plants by the development of seeds, and (2) vertebrates due to their evolution of the amniote
egg. Invasions of the aerial environment induced major radiations in insects about 325 Ma; pterosaurs by 251
Ma; birds by 150 Ma (Archaeopteryx or before); and bats by 52 Ma (but perhaps earlier). Terrestrial animals
and plants returning to the sea also caused rapid radiations: several clades of Mesozoic reptiles arose about 250
Ma, turtles by 150 Ma, mosasaurs by 85 Ma, mammals by 50 Ma, and sea snakes at 10 Ma.
Contemporary, invasion-caused adaptive radiations have been known to occur on both small and large
scales. Many small radiations have taken place in freshwater lakes or on terrestrial archipelagos. Extraordinary
assemblages of related species are often called “species flocks” and are known to occur among many different
animal groups. Some, such as the cichlid fishes of the African lakes, serve as textbook examples of rapid
speciation and diversification (Salzburger and Meyer, 2004). Other species flocks have been detected in
marine waters: the gastropod fauna of the Cape Verde archipelago contains 47 endemic and only three
nonendemic Conus species (Duda and Rolánd, 2005), rockfishes of the genus Sebastodes are concentrated in
the Northeast Pacific (Johns and Avise, 1998), and notothenioid fishes dominate the ichthyofauna of the
Antarctic (Eastman and McCune, 2000). Terrestrial species flocks are numerous including such well known
examples as those in the Hawaiian, West Indian, and Galapagos islands. Despite their limited geographic size,
these radiations have occurred over many parts of the world and produced endemic species that have made
significant additions to the global total. On an even smaller scale are the continuing invasions by individual
species. Over 300 species have invaded the Eastern Mediterranean via the Suez Canal (Galil, 2007), more than
200 have become established in San Francisco Bay (Ruiz et al., 1997), and many others are found in estuarine
and coastal environments around the world. Contemporary single invasions continuously add to local diversity
and, via speciation, have the potential of adding to the global total. All of the foregoing information, both
fossil and Recent, provides support to Simpson (1953) who thought that adaptive radiations could explain all
of life’s diversity, and to Schluter (2000) who observed that much of life’s diversity, perhaps even most of it,
arose from such episodes.

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8 Conclusions
Multiple invasions of animals and plants into contrasting habitats have occurred on a major and minor scale
from the early Paleozoic to Recent time, and have made notable contributions to global biodiversity. The land
and aerial invasions of the insects, by themselves, accounts for most of the world’s terrestrial species diversity.
However, the major radiations were not all confined to ancient times. The Cenozoic era, which began 65 Ma,
has seen evolution of most modern birds (Neornithes), including the origin of about 9,000 species. Mammalian
invasion of the sea, including the origin of the whales, pinnipeds and sirenians, has taken place within the past
50 Myr. Evolution of our present terrestrial mammal diversity, involving numerous regional invasions, also
took place within the Cenozoic. The radiation of sea snakes began only 10 Ma.
Over the course of the Phanerozoic, there have been evident decreases in the magnitude of adaptive
radiations and the size of their contributions to global biodiversity. This indicates that, despite the setbacks
caused by the five major extinctions, radiations became more restricted as ecological opportunity became less.
Even so, species invasions leading to biological innovation still occur and new habitats caused by tectonic and
climatic changes will continue to become available. Depending on size and complexity, the new habitats will
offer platforms for large or small adaptive radiations.
In our dynamic world, species are continually migrating, most commonly from high diversity areas to
those of lesser diversity. Species that successfully invade (colonize), add to local diversity. Over time, many of
the invaders will speciate, thus increasing global diversity. If we combine the diversity gains produced by the
three invasive catagories, i.e., global adaptive radiations, local radiations, and single species effects, that total
may account for much of the biodiversity gains that have taken place in the intervals between mass extinctions.
The present high level of the world’s biodiversity had its beginning with the advent of the CTR which took
place 125-80 Ma. The CTR was fueled by a huge increase in primary production provided by angiosperm
plants as they invaded the temperate zones of the Earth. The impetus from the CTR continued through the
Cenozoic despite a temporary setback caused by the K/T extinction. If we value and protect the individual
species that comprise the total biodiversity, it should continue to rise until the next mass extinction. A small
minority of invasive species are pests to human endeavors and need to be controlled, but for the sake of
biodiversity conservation, it would behoove us to judge each species on its own merits.
The positive contributions of historic and contemporary invasions to global biodiversity may be contrasted
to the claims of biodiversity loss that dominate the current literature. These claims have little validity and
species invasions, combined with ongoing allopatric, parapatric, and sympatric speciation, are continually
adding to biodiversity. The world’s primary conservation problem is not the loss of biodiversity, it is the loss
of habitat and over exploitation through human activity that has reduced thousands of formerly widespread
species to small remnant populations. Many of these vulnerable populations can still be rescued if there is
sufficient interest in doing so.

Acknowledgments
I wish to thank E.A. Hanni, M.K. Chew, A.L. Stigall and M. Dawson for discussion and helpful comments.

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Environmental Skeptics and Critics, 2014, 3(1): 17-23

Article

True-to-life friction values in connectivity ecology: Introducing

reverse flow connectivity

Alessandro Ferrarini
Department of Evolutionary and Functional Biology, University of Parma, Via G. Saragat 4, I- 43100 Parma, Italy
E-mail: sgtpm@libero.it,alessandro.ferrarini@unipr.it

Received 9 November 2013; Accepted 15 December 2013; Published online 1 March 2014

Abstract
Modelling ecological connectivity across landscape is pivotal for understanding a large number of ecological
processes, and for achieving environmental management goals such as preserving plant and animal populations,
predicting infective disease spread and conserving biodiversity. A pivotal topic in connectivity ecology is how
to assign realistic resistance (frictional) values to landscape categories. Based on these values and on the
dispersal model, a large number of ecological processes can be understood. While in a recent paper I proposed
a connectivity model and conceptual framework (flow connectivity) that is alternative to circuit theory, in this
paper I propose an advance to flow connectivity that is able to solve this topic. Thanks to the conceptual and
operative framework proposed here, named reverse flow connectivity, the subjectivity in ecological
connectivity is minimized. Since connectivity is a pillar of metapopulation theory and gene flow, reverse flow
connectivity can be regarded as a contribution to different fields of ecology, biology and landscape genetics.
Reverse flow connectivity can also be applied to conservation planning and for predicting ecological and
genetic effects of spatial heterogeneity and landscape changes.

Keywords flow connectivity; gene flow; landscape connectivity; metapopulation theory; partial differential
equations; resistance values; species dispersal.

Environmental Skeptics and Critics     
ISSN 2224­4263   
URL: http://www.iaees.org/publications/journals/environsc/online­version.asp 
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E­mail: environsc@iaees.org 
Editor­in­Chief: WenJun Zhang 
Publisher: International Academy of Ecology and Environmental Sciences 

1 Introduction
Modelling ecological connectivity across landscape is pivotal for understanding a large number of ecological
processes, and for achieving environmental management aims such as preserving plant and animal populations,
predicting infectious disease spread, and conserving biodiversity (Crooks et al., 2006; McRae, 2006).
Long-term persistence and stability of meta-populations within fragmented landscapes mainly depends on
re-colonisation between habitat patches (Gustafson and Gardner, 1996). Colonisation probability is implicitly
limited by inter-patch distance and dispersal ability of the species under study. Consequently, dispersal is the
key-process responsible for metapopulation persistence (Hanski, 1994; Moilanen and Hanski, 1998; Moilanen

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and Nieminen, 2002), determining the long-term survival of metapopulations due to demographic and genetic
effects (Taylor, 1990; Cohen and Levin, 1991; Poethke et al., 2003). Additionally, dispersal is an important
factor determining local extinctions (Poethke et al., 1996).
Understanding broad-scale ecological processes that depend on connectivity and incorporating such
connectivity into conservation planning needs to assess how connectivity is affected by environmental features
(McRae et al. 2007; McRae et al., 2008). Dispersal is affected by the quality of the matrix in which dispersal
happens (Gustafson and Gardner, 1996; Moilanen and Hanski, 1998; Wiegand et al., 1999; Wiens, 2001) and
by the distribution of suitable patches in the landscape (Zollner and Lima, 1999; King and With, 2002). There
is hence a demand for efficient and reliable models that relate landscape composition and pattern to the
connectivity of ecological processes. Recently, concepts and algorithms from electrical circuit theory have
been adjusted for these purposes (McRae, 2006; McRae et al., 2008). In circuit theory, landscapes are
represented as conductive surfaces, with resistance proportional to the easiness of species dispersal or gene
flow. Low resistances are assigned to habitats that are most permeable to movement or best boost gene flow,
and high resistances are given to poor dispersal habitat or to barriers. Circuit theory offers several advantages,
including a theoretical basis in random walk theory and the ability to evaluate contributions of multiple
dispersal pathways. For example, effective resistances calculated across landscapes have been shown to
markedly improve predictions of gene flow for plant and animal species. More details can be found in McRae
(2006), McRae (2007) and McRae et al. (2008).
Recently I introduced a modelling approach and theoretical framework named “flow connectivity”
(Ferrarini, 2013), that is alternative to circuit theory, and applied it as an example to the Ceno Valley (Ferrarini,
2005; Ferrarini et al., 2010; Ferrarini and Tomaselli, 2010; Ferrarini, 2011; Ferrarini, 2012a; Ferrarini, 2012b).
It is able to fix the weak point of the “from-to” connectivity approach, and it holds also for mountain and hill
landscapes. In addition, it doesn’t assume any intention for a species to go from source points to sink ones,
because the expected path for the species is determined locally (pixel by pixel, greedy model) by landscape
features.
By the way, a drawback is still open, i.e. how to assign true-to-life friction (resistance) values to landscape
categories. To this reason, I offer here a modelling solution to friction values assignment in connectivity
ecology. I’ve called this approach “reverse flow connectivity” since it makes use of flow connectivity
modelling (Ferrarini 2013), but it reverses the goal: instead of predicting species dispersal over landscape, it
builds up a proper friction landscape so that predicted and observed biotic flows coincide.

2 Reverse Flow Connectivity: Mathematical Formulation

Let L ( x, y , z , t ) be a real 3D landscape at generic time t, where L  [1,..., n] . In other words, L is a generic
(categorical) landcover or land-use map with n classes.
Let  ( L ) be the landscape friction (i.e. how much each land parcel is unfavourable) to the species under
study. In other words,  ( L) is a function that associates a friction value to each pixel of L.
Landscape friction has 2 components, i.e. the structural and the functional one, and the overall friction
should be equal to their product (not the sum) since they’re interactive:

 ( L )   STR ( L ) *  FUNC ( L ) (1)

Let Ls ( x, y ,  ( L )) be a landscape where, for each pixel, the z-value is equal to the friction for the species
under study. In other words, Ls is a 3D fictional landscape with the same coordinates and geographic
projection as L, but with pixel-by-pixel friction values in place of real z-values. Higher elevations represents
areas with elevated friction to the species due to whatever reason (unsuitable landcover, human disturbance

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 Environmental Skeptics and Critics, 2014, 3(1): 17-23 19

etc…), while lower altitudes represent the opposite. Barriers (structural or functional) can be simulated using
very elevated friction values.
Let S ( x , y , t ) be a binary landscape with the same coordinates and geographic projection as Ls and L,
but with binary values at each pixel representing species presence/absence at generic time t.
At time T0,

S 0  S ( x , y , t0 ) (2)

Now I have simulated the biotic flow over the fictional landscape Ls as follows (Ferrarini, 2013)

 S ( x, y , t ) S S
 div S    S   (3)
t x y
with initial conditions S0 at time T0.
As defined in Ferrarini (2013), the resulting biotic flow is as follows:


0 if  S   S  0
 x y


S  S S
 1 if (  1 and  0)
t  x y (4)

 S S
 or (  0 and  1)
 x y
 S S
 or  1
 x y
Fig. 1 illustrates the above-described concepts.

Fig. 1 The fictional landscape Ls has been built upon the real landscape L(x,y,z,t). Elevation represents for each pixel the
landscape friction  ( L ) for the species under study: the higher the elevation, the higher the friction to the species. Black points
represent sites where the species has been simulated to be present (S0=1) in Ferrarini (2013).

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20 Environmental Skeptics and Critics, 2014, 3(1): 17-23

My model assumes that the species dispersal ends at a stability point, if exists, where:

 S ( x, y , t )
 S  0 (5)
t
Now, I define P as the predicted path for the species over the fictional landscape Ls, and P* the real path
followed by the species as detected by GPS data-loggers or in situ observations. The prediction bias B between
P and P* can hence be calculated as (Fig. 2)

B  mod(  Pdx   P * dx ) (6)

where the function mod indicates the module of the difference, hence:

 Pdx  P*dx
  where P >P*
B (7)
  P dx   Pdx
*
where P* >P

Fig. 2 Expected biotic dispersal path P from simulated presence (black point) is depicted in red. The path ends when the model
detects a stability point. The path in magenta represents the detected path P* via GPS data-loggers. The area between the two
curves represents the prediction bias B. Both the predicted and the detected paths can be seen as curves in a XY-plane. Hence, the
bias B is equal to the area between the two curves. Reverse flow connectivity aims at setting B to 0 by optimizing friction
(resistance) values of land cover types.

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 Environmental Skeptics and Critics, 2014, 3(1): 17-23 21

Now, reverse flow connectivity acts as follows:

set B to 0 (8)
or, at least,
minimize B (9)

by optimizing  ( L) under the constraints that

 ( Li )  0 i
 (10)
 ( Li )  n i, with n integer value
In other words, reverse flow connectivity assigns realistic resistance values to each land cover type by making
null the bias B between the predicted dispersal and the detected one. To do this, it builds up the fictional 3D
landscape Ls ( x, y ,  ( L )) so that the predicted biotic flow P corresponds to the one (i.e. P*) detected in situ.
The optimization of  ( L) can be properly achieved using genetic algorithms (GAs; Holland, 1975). In
order to apply reverse flow connectivity modelling to real landscapes, I wrote the ad hoc software Reverse
Connectivity Lab (Ferrarini, 2013b). GAs are powerful evolutionary models with wide potential applications
in ecology and biology, such as optimization of protected areas (Parolo et al., 2009), optimal sampling
(Ferrarini, 2012c; Ferrarini, 2012d) and networks control (Ferrarini, 2011b; Ferrarini, 2013c; Ferrarini, 2013d;
Ferrarini, 2013e).
There are two useful suggestions when seeking true-to-life values for landscape resistance values. First, it
is always better to start with a 3D fictional landscape Ls ( x, y ,  ( L )) built upon friction values taken from
literature. In this way, it’s likely that reverse flow connectivity might need just minor corrections to such
values when optimizing  ( L) in order to set B to 0. Instead, if the researcher starts from completely
unknown values for  ( L) (e.g. friction values set to 0 for each land category), it is possible that the
computational effort becomes very high for common computers. Of course, the computational effort also
depends on the number of land categories and on the length of the path P*. Secondly, a potential drawback
arises when the real path P* does not meet all the land cover categories present in the landscape. In this case,
reverse flow modelling will be only able to assign resistance values to those land categories that have been
intersected by P*.

3 Conclusions
The pivotal topic in connectivity ecology is how to assign realistic resistance (frictional) values to landscape
categories. Based on these values and on the dispersal model, a large number of ecological processes can be
understood, and many environmental management goals can be achieved such as preserving plant and animal
populations, predicting infective disease spread and conserving biodiversity.
While in a previous paper (Ferrarini, 2013) I proposed a connectivity model and conceptual framework that
is alternative to circuit theory, in this paper I have proposed an advance to flow connectivity that is able to
assign true-to-life frictional values to landscape categories. Thanks to the conceptual and operative framework
proposed here, named reverse flow connectivity, the subjectivity in ecological connectivity is minimized and
proper prediction of population dispersals across landscape is now possible.
Since connectivity is a pillar of metapopulation theory and gene flow, reverse flow connectivity can be
regarded as a contribution to different fields such as ecology, biology and landscape genetics. Reverse flow
connectivity can also be applied to conservation planning and for predicting ecological and genetic effects of
spatial heterogeneity and landscape changes.

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22 Environmental Skeptics and Critics, 2014, 3(1): 17-23

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Environmental Skeptics and Critics
ISSN 2224-4263
Volume 3, Number 1, 1 March 2014

Articles

First evidences of sexual divergences in flight behaviour and space use of lesser kestrel
Falco naumanni
Marco Gustin, Alessandro Ferrarini, Giuseppe Giglio, et al. 1-7

Invasions, adaptive radiations, and the generation of biodiversity

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True-to-life friction values in connectivity ecology: Introducing reverse flow connectivity

Alessandro Ferrarini 17-23

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