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Seasonality of reproduction is observed in many species of organisms, across taxa, and is influenced by both biotic and abiotic factors. While such seasonality is easy to understand in temperate species exposed to extreme climates, it is more difficult to explain in the tropics. In many tropical species offspring are born during the season of high precipitation, which also coincides with high resource availability. Interestingly, in India, free-ranging dogs seem to mate, and not whelp, when it rains-an observation that cannot be explained by the resource abundance hypothesis. We carried out an extensive study to identify the mating seasons of free-ranging dogs, and observed a strong correlation between both the incidence and frequency of mating related behaviours of dogs, and precipitation levels. There are two clear mating seasons, of which the primary mating season coincides with the monsoon (rainy season) and the secondary mating season coincides with the nor'westerlies in this part of India. We speculate that this strong correlation is an effect of chemistry, rather than biology. While male dogs can mate round the year, females come into estrous seasonally. In the urban environment, dogs are exposed to a lot of olfactory noise, which can dilute the signal present in sex pheromones of the females in heat. A shower leads to increased humidity and reduced temperature of the air, leading to intensification of pheromone signals that trigger a sexual response in the dogs.
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When Love Is in the Air: Understanding Why
Dogs Tend to Mate when It Rains
Sreejani Sen Majumder, Anindita Bhadra*
Behaviour and Ecology Lab, Department of Biological Sciences, Indian Institute of Science Education and
ResearchKolkata, Kolkata, India
Seasonality of reproduction is observed in many species of organisms, across taxa, and is
influenced by both biotic and abiotic factors. While such seasonality is easy to understand
in temperate species exposed to extreme climates, it is more difficult to explain in the trop-
ics. In many tropical species offspring are born during the season of high precipitation,
which also coincides with high resource availability. Interestingly, in India, free-ranging
dogs seem to mate, and not whelp, when it rainsan observation that cannot be explained
by the resource abundance hypothesis. We carried out an extensive study to identify the
mating seasons of free-ranging dogs, and observed a strong correlation between both the
incidence and frequency of mating related behaviours of dogs, and precipitation levels.
There are two clear mating seasons, of which the primary mating season coincides with the
monsoon (rainy season) and the secondary mating season coincides with the norwesterlies
in this part of India. We speculate that this strong correlation is an effect of chemistry, rather
than biology. While male dogs can mate round the year, females come into estrous season-
ally. In the urban environment, dogs are exposed to a lot of olfactory noise, which can dilute
the signal present in sex pheromones of the females in heat. A shower leads to increased
humidity and reduced temperature of the air, leading to intensification of pheromone signals
that trigger a sexual response in the dogs.
Most species of mammals exhibit some degree of reproductive synchrony, so that breeding
occurs during optimal conditions [1,2]. Seasonal breeders successfully mate only during a par-
ticular time of the year, typically giving birth to offspring at a time which is ideal for the sur-
vival of the young [3]. Seasonality of reproduction is observed in mammals across all latitudes
and has even been suggested in the archetypal example of continuously breeding species,
Homo sapiens [4]. This can be observed in animals, both in terms of the onset of reproductive
maturity for young individuals, as well as reproductively active phases for the population in an
annual scale [5]. Animals living in seasonally harsh climates tend to display sharply delineated,
short breeding seasons, while those that live in relatively non-varying environments display
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 1/15
Citation: Sen Majumder S, Bhadra A (2015) When
Love Is in the Air: Understanding Why Dogs Tend to
Mate when It Rains. PLoS ONE 10(12): e0143501.
Editor: Hiroaki Matsunami, Duke University, UNITED
Received: July 13, 2015
Accepted: November 5, 2015
Published: December 2, 2015
Copyright: © 2015 Sen Majumder, Bhadra. This is
an open access article distributed under the terms of
the Creative Commons Attribution License, which
permits unrestricted use, distribution, and
reproduction in any medium, provided the original
author and source are credited.
Data Availability Statement: All data are available in
the paper. The authors have provided the raw data
and relevant analysis in the supporting information
Funding: The work was funded by the Women's
Excellence Award No. SB/WEA-005/2013 by the
Science and Engineering Board, Department of
Science and Technology, India to AB; Indian Institute
of Science Education and Research Kolkata, India
provided infrastructural support.
Competing Interests: The authors have declared
that no competing interests exist.
little or no seasonality of reproduction [1]. Seasonality of reproduction can be strongly influ-
enced by abiotic factors like variations of day length, temperature and humidity, which in turn
can affect resource availability [613]. In fact, it has been shown, at least in the case of primates,
that the effects of climate and latitude on birth seasonality are mediated via the availability of
resources. Temperate species are more sensitive to changes in day length and temperature,
while tropical species are more affected by rainfall patterns. The width of the peak of birth sea-
sonality can be extremely variable, and is often dependent on multiple environmental as well as
biological factors [14].
Why do animals in the tropics display seasonality of reproduction? It has been suggested
that seasonal breeders are able to maximize fitness in individuals by synchronizing energeti-
cally demanding periods of the breeding cycle with periods of maximum food availability or
quality, by giving birth just before or during the peak in resource availability [5,1521]. Lacta-
tion is the most energetically costly phase in the reproductive cycle of female mammals; it is
thus beneficial to the female to match this phase with a period of resource abundance, so that
she has access to food supply to replenish her lost reserves of energy [1]. This idea is substanti-
ated by observations on many species like giraffes (Giraffa camelopardalis), African elephants
(Loxodonta africana), gazelles, rhesus macaques (Macaca mulatta) and wolves (Canis lupus)
[2227]. The fact that seasonal breeders can become continuous breeders in resource abundant
conditions like domestication and in zoos also lends support to this hypothesis [1,28,29].
Though seasonality of reproduction is well understood in temperate species, especially with
respect to shift in day length, the mechanism of seasonality in tropical animals is open to explo-
ration. The most accepted hypothesis to explain seasonality of reproduction in the tropics per-
tains to the maximum availability of resources during the wet season [3034]. Is the availability
of resources the only cause of increased reproductive activities during the wet season in the
tropical regions? More importantly, how does rainfall trigger reproductive activity in the
Dogs (Canis lupus familiaris) are the first species to have been domesticated [29], and
though most canids are known to be seasonal breeders [27,29,3537] domestic dogs are
known to breed continuously, with no clear mating season [3840]. However, indirect evidence
for seasonality of breeding have been suggested [41], especially in free-ranging dogs [42]. In
India, free-ranging domestic dogs are a ubiquitous presence in all kinds of human habitats,
leading a life of scavengers [43,44]. It has been observed that the free-ranging dogs in West
Bengal, India have a clear mating season, which coincides with the monsoon or the wet season
[45,46]. Monsoon is also the mating season for various other mammals in the Indian subconti-
nent [30,47,48]. This presents a very intriguing paradoxmating, and not whelping, occurs
during the monsoon, with the offspring mostly being born during the winter, and so cannot be
explained by the hypothesis of resource abundance. The case of the dogs is all the more inter-
esting because being scavengers mostly dependent on wastes and offerings from humans for
their sustenance [44,49], their resources are expected to be constant throughout the year.
However, while pet dogs can reproduce aseasonally [50], free-ranging dogs seem to show defi-
nite seasonality of reproduction, mating during the monsoon and whelping in the winter. Thus
the rains somehow trigger reproductive activities, and this does not apparently provide any
adaptive advantage to the breeders.
In this paper we show that not only is there a strong concurrence between precipitation and
the mating behaviour of free-ranging dogs, but mating is highly correlated with precipitation
levels. We would like to suggest that the proximate cause for this strong correlation lies in abi-
otic, rather than biotic mechanisms, leading to the seasonal mating habit observed in free-rang-
ing dogs.
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 2/15
The Animal Ethics Committee, Indian Institute of Science Education and Research Kolkata
provided permission for this work.
The work reported here included two different sampling exercises, one involved a year-long
population level census, while the other involved observations of free-ranging dog groups dur-
ing the monsoon for four years.
Long term data
We carried out behavioural observations on free-ranging dogs in Kolkata (22.5667° N,
88.3667° E), West Bengal, India from the year 2010 to 2013. The study was conducted from
July to 15
October for the first three years, and from 15
June to 15
October for 2013.
The period of the study was based on earlier observations of mating during the monsoon and
pup occurrence in the winter [46,5153]. In West Bengal, monsoon arrives around mid-June,
with the thrust beginning in July; monsoon recedes in SeptemberOctober [54]. A neighbour-
hood in Saltlake, Kolkata was chosen for the observations, based on convenience and safety of
sampling during late hours. Dogs were observed three days a week from 17:00h to 00:00h using
instantaneous scans and all occurrences sampling. Each dog group was observed for two hours
every day from this above mentioned time period randomly, so that we got 6 hours of data for
each dog group per week. We collected data for a total of 954 hours on 67 free-ranging dogs
belonging to 12 groups, over 159 days. The area of this study was constant over the four years,
but the dogs varied between the years due to natural fluctuations in the population. The precip-
itation and temperature for each day of observation was recorded from the website of the India
Meteorological Department ( At the time of observations, we also
made a qualitative note of the weather conditions, like dry, mild drizzle, medium rainfall or
heavy rainfall. This qualitative record was later used to quantify precipitation levels into three
categorieslow (drylight drizzle), medium (short intense shower and light but steady rain
over 23 hours), high (heavy and prolonged rain). Table 1 provides an ethogram of behaviours
used in this study (also see S1 Fig).
Year-long census
We selected 40 locations randomly from Kolkata (22.5667° N, 88.3667° E), Kalyani (22.9750°
N, 88.4344° E), Kanchrapara (22.9700° N, 88.4300° E), Barrackpore (22.7600° N, 88.3700°E)
and Barasat (22.7200° N, 88.4800° E), West Bengal, India covering rural, urban and semi urban
areas (S2 Fig). A census of free-ranging dogs was carried out in these locations, covering each
of the 40 locations once over a three month period for a year (15
April 201414
2015). Thus we collected data from 510 locations every fortnight, covering each of the 40 loca-
tions four times during the year. Each three-month period overlapped, but did not completely
coincide with, a season in West Bengal [55]summer (mid-Aprilmid-July), monsoon (mid-
Table 1. Ethogram of mating related behaviours (MRB) used in the study.
Behaviour Description
Genital snifng (GS) A male sniffs at the genitalia of the female
Try to clasp (TC) A male tries to clasp a female from the back
Marking with urine (MK) Scent marking by leg lifting
Running together (RT) Running together male and female
Mount (MT) A male climbs over a female in an attempt to mate; usually leads to copulation
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 3/15
Julymid-October), winter (mid-Octobermid-January), spring (mid-Januarymid-April).
The time of census was fixed between 0600-0900h and 1600-1900h when dogs are typically
seen to be active on streets. We thus had 160 censuses in the whole year from the 40 locations,
and an additional set from 5 of the locations within a fortnight (1530 April 2015) so that the
sampling period could be circular for part of the analysis. For the censuses we followed the spot
sampling method described in [51]. The observer walked along all roads and by lanes of each
location and whenever a dog was sighted, the age category (Adult/Juvenile/Pups) of the dog,
its group size, sex and the behaviour at the time of sighting were recorded. The weather condi-
tion at the time of the census was recorded as described above, and the temperature and precip-
itation levels for the day were also recorded from the IMD website. We divided the behaviours
into two categories, mating-related and non-mating behaviours. Table 1 provides an ethogram
of all the mating-related behaviours observed.
Statistical Analysis
Using the actual levels of precipitation recorded on the day of observations (from IMD), a
regression analysis was carried out using data pooled across years to test if the frequency of
MRB depended on the precipitation levels. Repeated measures ANOVA were used to test for
variation in MRB between years, and between weeks of observation (within a year). We divided
the precipitation levels on the days of observation into three categories: high (>20mm per
day), medium (1020 mm per day) and low (010 mm per day). Frequency of MRB at different
precipitation levels were compared using a Kruskal-Wallis test. A full factorial ANOVA with
post hoc Tukeys test was conducted for the entire dataset, considering the frequency of MRB
as the dependent variable, and years, weeks, precipitation levels and behaviours as independent
variables. ANOVA with post hoc tests for different levels of precipitation over 12 weeks of 4
years was used to understand the variations at the level of each mating related behaviour. In
the year-long census, a repeated measures ANOVA was used to check for variations in dog
numbers between seasons in the 40 locations. Kruskal-Wallis tests were carried out to test for
the effect of season on the occurrence of MRB. All statistical analysis was carried out in Statis-
tiXL 2007. A distribution fitting exercise was carried out in the software Igor Pro using the fre-
quency of MRB and precipitation data for every fortnight.
Long term data
Mating related behaviours (MRB) were observed in all four years during the monsoon [56]. In
2013, MRB were first observed in the month of June, while in all the other years the onset of
mating was in the month of July. The occurrence of MRB reduced after the month of Septem-
ber. Hence observations spanned from June to September for 2013, and from JulySeptember
for the other three years. The frequency of MRB was strongly dependent on the precipitation
levels on a given day (Linear Regression: R
= 0.612, F = 223.694, p <0.001; Fig 1). There was
significant variation in the frequency of MRB across the years when the data for the period July
September was considered (Repeated measures ANOVA, F
= 7.696, p <0.0001). Post
hoc analysis revealed that there was significant difference in the frequency of MRB between
2013 and the other three years, while there was no difference in the levels of MRB between
2010, 2011 and 2012 (Table 2). Interestingly, when the frequency of MRB for first three months
after the onset of mating were considered, i.e., June to August for 2013 and JulySeptember
for the other years, there was no significant difference between the years within the same pre-
cipitation category (Repeated measures ANOVA, F
= 1.670, p = 0.187). This suggests that
the mating behaviour in dogs has some pattern that might be correlated with the precipitation
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 4/15
pattern. In both the above cases, there was no significant variation in MRB within a precipita-
tion category when the data was considered at a weekly level (Repeated measures ANOVA,
= 1.151, p = 0.353; F
= 1.472, p = 0.185). For all subsequent analysis, the data for the
JuneAugust period of 2013. i.e. for the first three months after the onset of mating was
The frequency of MRB was highest for the high precipitationcategory2703 acts of MRB
were observed over the four years on days that received high precipitation, while this number
was 778 and 252 in the medium and low precipitation categories. The variation in the fre-
quency of MRB for different precipitation categories was significant (Kruskal-Wallis test: χ
8.540, df = 2, p = 0.014). The full factorial ANOVA was highly significant (F
= 43.082,
p<0.0001), with only the interaction between year and precipitation level being non-signifi-
cant (Table 3). Post hoc tests revealed significant differences between all behaviours, and
between all the three precipitation levels. But variation within the years was not significant (Fig
2,Table 3).
Since there was significant variation between the behaviours, an ANOVA with post hoc
tests at the level of each behaviour was used to test for variations for occurrence of the behav-
iour at different levels of precipitation. For all behaviours other than MT, the frequency of
occurrence of the behaviour was significantly different between all three precipitation levels,
while the rate of MT was different only for the high precipitation category (S1 Table;Fig 3).
Fig 1. The occurrence of mating related behaviours (MRB) was highly correlated with precipitation levels (mm). The precipitation levels reported here
are actual readings of precipitation on the day of observations, as given by IMD. The linear fit is given by the black line, represented by the equation
y = 0.4888x + 18.893, R
= 0.6117.
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 5/15
Year-long census
We sampled an average of 15 dogs per location, considering all 40 locations for four seasons,
where the dog numbers varied from 559 (monsoon) to 645 (spring) for the 40 locations taken
together (S1 and S2 Figs). There was significant variation in dog numbers between seasons
(Repeated measures ANOVA: F
= 2.793, p = 0.003). However, the sex ratio did not deviate
from 1:1 across the seasons (T test for each season, p <0.05 for each season). We carried out a
Kruskal-Wallis test for the frequency of MRB observed in the four seasons over the six fort-
nights in each season. There was significant variation between seasons, when all four seasons
Table 2. Results of the post hoc Tukeys test for Repeated Measures ANOVA carried out to check whether there was significant variation between
the years (20102013) for the occurrence of mating related behaviours at different levels of precipitation (in mm) for three months from the onset
of mating activities (July to September for the first three years, June to August for 2013).
Test Y Variable Group 1 Group 2 Mean Diff. SE q Probability
Tukey >20 mm 2010 2011 6.500 4.446 1.462 0.731
2012 -2.417 4.446 0.544 0.980
2013 -4.000 4.446 0.900 0.920
2011 2012 -8.917 4.446 2.005 0.495
2013 -10.500 4.446 2.361 0.352
2012 2013 -1.583 4.446 0.356 0.994
1020 mm 2010 2011 4.500 2.132 2.110 0.451
2012 1.833 2.132 0.860 0.929
2013 4.667 2.132 2.188 0.419
2011 2012 -2.667 2.132 1.251 0.813
2013 0.167 2.132 0.078 1.000
2012 2013 2.833 2.132 1.329 0.784
nill 2010 2011 0.333 1.466 0.227 0.998
2012 -2.833 1.466 1.933 0.527
2013 -0.583 1.466 0.398 0.992
2011 2012 -3.167 1.466 2.160 0.430
2013 0.250 1.466 0.171 0.999
2012 2013 3.417 1.466 2.331 0.363
Table 3. Results of a Full Factorial ANOVA to test for interaction between precipitation levels, occurrence of MRB and years of observation.
Overall test of model for Y = Frequency
Source Type III SS Df Mean Sq. F Probability
Model 27666.682 59 468.927 43.082 0.000
Error 7183.750 660 10.884
Total 34850.432 719
Tests of effects for Y = Frequency
Source Type III SS Df Mean Sq. F Probability
Behaviour 8288.758 4 2072.190 190.380 0.000
Precipitation 13874.586 2 6937.293 637.357 0.000
Year 92.960 3 30.987 2.847 0.037
Behaviour*Precipitation 4117.275 8 514.659 47.284 0.000
Behaviour*Year 626.019 12 52.168 4.793 0.000
Precipitation*Year 102.769 6 17.128 1.574 0.152
Behaviour*Precipitation*Year 564.314 24 23.513 2.160 0.001
When Love Is in the Air
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were considered (χ
= 14.172, df = 3, p = 0.003). The significant variation was explained by the
increased rate of mating related activities in the monsoon, as the significance disappeared
when the three other seasons were compared (χ
= 2.984, df = 2, p = 0.225), but was evident for
all comparisons involving the monsoon (Fig 4).
The frequency of MRB observed per location within a fortnight strongly depended on the
average precipitation level in that fortnight (Linear regression: R
= 0.470, F = 20.428,
p<0.0001; Fig 5). The fortnightly data for precipitation levels and frequency of observed MRB
fitted double normal distributions, and there was an offset in both the sets of peaks for the two
distributions (Fig 6). This confirmed unambiguously the close correlation between precipita-
tion levels and mating activities of dogs.
Fig 2. The frequency of MRB varied with precipitation levels. The bar chart shows the mean and standard deviation of the frequency of all MRB occurring
at different levels of precipitation (as noted during the time of observations), over four years, 2010 to 2013, during the primary mating season. Variation in the
frequency of MRB was not significant across the four years within a precipitation category (alphabets), but varied significantly different levels of precipitation
within a year (*).
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 7/15
Reproductive activity in free-ranging dogs of West Bengal is not only seasonal, but shows spec-
tacular concordance with precipitation levels. Our results show a strong correlation between
precipitation levels and the occurrence of mating related behaviours in dogs, leading to a pri-
mary mating season during the monsoon and a secondary mating season that coincides with
the Norwesters or Kal Baisakhisin West Bengal [55], as revealed by our year-long census.
While the census based study revealed the strong connection between mating activities of dogs
and rainfall, the long term study of mating during the monsoon showed that there is a high
degree of consistency in the mating activities over the four years of the study, at least during
the primary mating season. It was interesting to note that the only deviation occurred when we
compared our observation for 2013 with the remaining data, keeping the calendar month con-
stant and ignoring the actual onset of rains. This difference vanished when we considered the
data from the actual onset of rains, and was consistent with the fact that there was significant
variation in MRB between the different weeks of observations within the three-month period
of the monsoons.
In this study, we were interested in understanding the extent to which free-ranging dogs
show seasonality of mating, and attempting to explore the plausible explanations for this
behaviour. Our observations on free-ranging dogs suggest that their attempts at mating are
Fig 3. All MRB showed variation across precipitation categories. Mean and standard deviation of the frequency of different mating related behaviours
observed at the three precipitation levels noted during the time of observationsthe different alphabets represent significant differences within a behaviour
category, between precipitation levels.
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 8/15
often disrupted by people, mostly due to socio-cultural reasons, and this reduces the probabil-
ity of seeing successful ties on the streets. Hence we used all activities that pertain to mating in
the dogs, and not just ties, but did not use mating related aggression for this analysis. We used
multivariate statistics to gain a more detailed understanding of the relationship between the
mating behavioural repertoire of the dogs and rainfall. Our observations revealed that all mat-
ing related behaviours are affected by rainfall, but the fact that mounting (MT) occurs at signif-
icantly higher rates at high precipitation levels only, suggests that precipitation levels might
eventually drive mating success. But why is mating in dogs influenced so intensely by precipita-
tion levels?
The answer to this question might lie in chemistry, rather than biology. Odor is a generic
term used for any compound or mixture of compounds that can be detected by animals
through olfaction. Though the word odor is usually associated with unpleasant smells, techni-
cally, even pleasant smells are odors. The ability to detect odors varies greatly among species,
and dogs are known for their highly developed olfactory acuity. They can detect odorant con-
centration levels at 12 parts per trillion, which makes them 10
times more efficient in
sensing odors than humans [57,58]. This fantastic sense of smell was probably one of the
major factors that made dogs useful to early humans as excellent hunting companions, and
also makes them indispensable in todays world. Sniffer dogs are used for tasks as diverse as
tracking criminals, locating survivors at sites of disasters, unearthing explosives and narcotics
and even estimating animal populations from scat samples [59].
The efficiency of dogs at tracking and locating is known to be influenced by environmental
factors like temperature and humidity. Tracking dogs are able to pick up trails better that are
laid on moist, rather than dry soil, and dogs trained for hunting birds are most efficient when
Fig 4. The year-long census showed that highest MRB occur in the monsoon. A bar chart showing the mean and standarddeviation of the frequency of
MRB per location in the four seasons. Different alphabets represent significant differences in the frequency of MRB between seasons. 40 locations were
used in the study.
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the ground is damp. Search-and-rescue dogs also perform better when the relative humidity is
high [60]. Humans as well as birds continuously slough off dead skin containing live bacteria.
The bacteria metabolize the dead cells and associated body secretions, leading to the formation
of odor, which is used by dogs for tracking. High temperature and dry conditions can kill bac-
teria and prevent odor formation, while very cold conditions retard bacterial action, and thus
decay rate. A light rain after a dry period can induce bacterial growth and hence the production
of odor. Thus humidity but not very high rains increase the efficiency of dogs in tracking and
searching tasks by increasing odor intensity [61]. Dogs also perform better at picking up trails
when the ground is warmer than the air, because the odor rises upwards in the cooler air [60].
Olfactory function in humans have also been shown to be affected by environmental condi-
tionsolfactory sensitivity thresholds are low in high humidity and low atmospheric pressure
Ion mobility spectrometry (IMS) is a widely used technique used for detecting trace quanti-
ties of gaseous organic compounds at atmospheric pressure, and is widely used for detecting
noxious elements [6366]. One major problem associated with this technique is the presence
of moisture, and currently a great deal of research focuses on dealing with this issue [67,68].
Humidity affects the detection process in IMS by reducing both selectivity and sensitivity [69],
as water interacts with ions to cause shift in peaks or the formation of additional peaks [70].
Odoriferous molecules also tend to interact with water vapour in the atmosphere, and this
slows down their diffusion in the air, and humidity reduces the threshold level of olfactory sen-
sitivity [62].
Fig 5. The frequency of MRB per location increased with increased precipitation. A scatterplot showing the significant interaction between the numbers
of MRB observed per location at different levels of precipitation (as recorded from IMD). The dotted line shows the linear fit, and is represented by the
equation y = 0.1243x + 0.581, R
= 0.4704.
When Love Is in the Air
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We speculate that the increased mating related activities of dogs during the monsoon is
brought about by a three-pronged effect. Firstly, heavy showers serve to wash away odors pres-
ent in the air so that there is less olfactory noiseimmediately after a shower. Due to the
increased humidity, the sex pheromones released by the individuals diffuse slowly, and thus
can be perceived in high concentration around the individuals. In contrast to the IMS instru-
ments, the dogs perform better under reduced temperature and increased humidity conditions,
which ensures that the higher concentration of sex pheromones in the air triggers higher mat-
ing related behaviours. The offset between peaks for highest precipitation MRB level probably
arises because extremely heavy rains can also wash away the sex pheromones, thereby reducing
their efficacy.
The absence of seasonality of reproduction in pet dogs [50] is likely to have arisen from the
availability of abundant resources [1]. Most canids show strong seasonality of reproduction in
the wild, the breeding season being typically determined by resource availability [27,35,37].
Since the free-ranging dogs are scavengers, the weather-induced seasonality of mating is likely
to increase competition for resources during the breeding season, rather than ensuring resource
abundance. It has been shown that male dogs get sexually excited on perceiving the urine and
vaginal discharge of females in heat [71], and they also sniff and lick the anogenital regions of
Fig 6. MRB and precipitation patterns show convergent double binomial trends. The plot shows the relationship between the occurrence of mating
related behaviours (MRB) and precipitation levels over a year, on a fortnightly basis. The red dots represent the number of MRB averaged over the numberof
locations sampled in a fortnight and blue dots represent the average precipitation level (from IMD)considering only the days of the census. Both the datasets
fit double normal distributions, as represented by the lines of the respective colours. A1 and A2 represent the two amplitudes of the respective curvesatx0
time as read from the x-axis.
When Love Is in the Air
PLOS ONE | DOI:10.1371/journal.pone.0143501 December 2, 2015 11 / 15
females in the proestrous stage [72]. Though male dogs find estrous urineto be an attractant,
such urine loses its attractiveness when stored for an hour at room temperature. This suggests
that the sex pheromone, if present, is not very stable, and is perhaps quite volatile [72]. It
should be noted that sexual activity in females also increases when they are in heat, leading to
increased investigation of male dogs and their urine [71]. Hence it seems likely that the
increased sexual activity of dogs immediately after a shower is triggered by the increased con-
centration of odorants in the humid air due to a slower rate of diffusion. This would be further
enhanced by the increased sensitivity to odor in humid conditions, for a short period of time
while the cues from the proestrous females persist. Though such weather-induced mating
behaviour can effectively increase the probability of mating success simply by increased mating
attempts, it could also be effectual as a natural population control mechanism of the free-rang-
ing dogs, due to the increase in competition for resources.
Supporting Information
S1 Fig. A map showing the position of the area where the long term behavioural observa-
tions on mating were carried out. The position of each group is marked with a yellow tag.
S2 Fig. A map showing the 40 locations in which the year-long census was conducted
pointed out using yellow tags.
S1 Table. A table giving the results of the ANOVAs conducted at the level of each mating
related behaviour for variation across years at different levels of precipitation.
The work reported in this manuscript was conceptualized by AB, SSM carried out the field
work under the supervision of AB. SSM and AB carried out the analysis and co-wrote the man-
uscript. The authors would like to thank Dr. Ayan Banerjee, Department of Physical Science,
Indian Institute of Science Education and Research Kolkata for his help with the curve fitting
exercise reported in Fig 3, and Ms. Manabi Paul, Department of Biological Sciences, Indian
Institute of Science Education and Research Kolkata for her help with editing the manuscript.
This work was funded by grant no. SB/WEA-005/2013 by the Science and Engineering
Research Board, Department of Science and Technology, Government of India to AB, and sup-
ported by the Indian Institute of Science Education and Research Kolkata. The institute animal
ethics committee approved of the research reported in this paper, and comply with the laws
pertaining to animal rights in India. We certify that the work was purely observation based,
and no animals were disturbed or harmed in any manner while carrying out the observations.
Author Contributions
Conceived and designed the experiments: AB. Performed the experiments: SSM. Analyzed the
data: SSM AB. Wrote the paper: SSM AB.
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... A recurrent dynamic revealed in HSI's street dog monitoring surveys is the seasonal fluctuation of street dog abundance when examined on a biannual survey cycle (1)-the result of apparent breeding seasonality in the street dog population. Elsewhere, seasonality of oestrus has been observed amongst free-roaming dog populations in the Indian settlements of Jodhpur (2), Jaipur (3), West Bengal (4), Katwa town (5), Kolkata (6), Goa and Tamil Nadu (7); however, other studies have asserted that this dynamic is not universal to free-roaming populations of the species (8), nor for many confined populations (9,10). Indeed, the reproductive cycle of the domestic dog (Canis familiaris) has long been considered non-seasonal (11,12). ...
... Thus, the question then becomes, what influences are the cause of seasonal breeding tendencies in the street dog populations discussed, and specifically, why post-monsoon? A number of factors have been identified, including increased food availability, artificially altered exercise and housing conditions, favorable climatic conditions, precipitation levels and sensitivity to daylight hours (3,6,7,9). ...
... Additionally, other seasonal cues may act to trigger seasonal reproductive dynamics independent of or in addition to photoperiod, particularly in regions of lesser correlation between food availability and day length, such as arid and equatorial environments (27). Such secondary triggers include rainfall (27), via its impact on pheromone dissipation for example (6), temperature (17,27), and other more species-specific triggers (26). In the case of the latitudes represented in this study, where little seasonal variation in photoperiod occurs, it may be that such secondary triggers are pertinent. ...
Full-text available
Understanding seasonal breeding dynamics is essential for maximizing efficiency and welfare in the application of street dog management programs. Humane Society International (HSI) has conducted many animal birth control programmes concerning the street dog populations in urban India. This paper analyses the data on reproductive indicators—oestrus, pregnancy, and pups—collected by HSI sterilization clinics in the cities of Jamshedpur, Dehradun, and Vadodara over a period of 3, 5, and 4 years, respectively. We found a consistent reproductive seasonality dynamic in all three cities, with peak oestrus and pregnancies occurring in the late/post-monsoon season. Pup proportions peaked soon after. Both these findings are consistent with previous studies of free-roaming domestic dog populations both in India and worldwide. Additionally, we identified minor inter-city differences in the temporal breeding dynamic, which we propose are the result of localized seasonal climatic and human factors. Finally, we examine and assert the relevance of breeding seasonality in the implementation of efficient and welfare-sensitive birth control programmes.
... Both in order to study free-ranging dogs and to compare their behavioural patterns with other canids, there is a need for an ethogram that is accessible to all. We have carried out detailed behavioural observations of freeranging dogs in their natural habitats in India for several years and published partial ethograms focusing on certain aspects of their behaviour, like foraging, parental care and mating (Sen Majumder et al. 2014a, 2014bSen Majumder and Bhadra 2015;. ...
The domestic dog (Canis lupus familiaris) is known to have evolved from gray wolves, about 15,000 years ago. They frequently exist as free-ranging populations across the world. They are typically scavengers and well adapted to living among humans. Most canids living in and around urban habitats tend to avoid humans and show crepuscular activity peaks. In this study, we carried out a detailed population-level survey on free-ranging dogs in West Bengal, India, to understand the activity patterns of free-ranging dogs in relation to human activity. Using 5669 sightings of dogs, over a period of 1 year, covering the 24 h of the day, we carried out an analysis of the time activity budget of free-ranging dogs to conclude that they are generalists in their habit. They remain active when humans are active. Their activity levels are affected significantly by age class and time of the day. In addition, we provide a detailed ethogram of free-ranging dogs. This, to our knowledge, is the first study of this kind, which might be used to further study the eco-ethology of these dogs.
... Moreover, according to Gutzwiller [44], increased humidity could be responsible for increased odor intensity, positively influencing the tracking efficiency of dogs. This phenomenon is also thought to be true in the context of semiochemical substance detection [45]. However, even though the increased humidity connected with light rain is usually perceived as a positive influence on odor detection, heavy rain is usually a negative factor, since it could force the scent down lower to the ground [46]. ...
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Olfaction in dogs is crucial for gathering important information about the environment, recognizing individuals, making decisions, and learning. It is far more specialized and sensitive than humans’ sense of smell. Using the strength of dogs’ sense of smell, humans work with dogs for the recognition of different odors, with a precision far exceeding the analytical capabilities of most modern instruments. Due to their extremely sensitive sense of smell, dogs could be used as modern, super-sensitive mobile area scanners, detecting specific chemical signals in real time in various environments outside the laboratory, and then tracking the odor of dynamic targets to their source, also in crowded places. Recent studies show that dogs can detect not only specific scents of drugs or explosives, but also changes in emotions as well as in human cell metabolism during various illnesses, including COVID-19 infection. Here, we provide an overview of canine olfaction, discussing aspects connected with anatomy, physiology, behavioral aspects of sniffing, and factors influencing the olfactory abilities of the domestic dog (Canis familiaris).
... Secondly, the environmental conditions influence the odour sensing abilities. Relative humidity and barometric pressure may directly affect olfaction, besides their effects on odour emergence and movement itself, while heat has only indirect effects [58,[82][83][84]. Acclimatization to the environment, physical fitness and an adequate hydration state can prevent heat stress of the dogs [84]. ...
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The extraordinary olfactory sense of canines combined with the possibility to learn by operant conditioning enables dogs for their use in medical detection in a wide range of applications. Research on the ability of medical detection dogs for the identification of individuals with infectious or non-infectious diseases has been promising, but compared to the well-established and–accepted use of sniffer dogs by the police, army and customs for substances such as money, explosives or drugs, the deployment of medical detection dogs is still in its infancy. There are several factors to be considered for standardisation prior to deployment of canine scent detection dogs. Individual odours in disease consist of different volatile organic molecules that differ in magnitude, volatility and concentration. Olfaction can be influenced by various parameters like genetics, environmental conditions, age, hydration, nutrition, microbiome, conditioning, training, management factors, diseases and pharmaceuticals. This review discusses current knowledge on the function and importance of canines’ olfaction and evaluates its limitations and the potential role of the dog as a biomedical detector for infectious and non-infectious diseases.
... The fact that free-ranging dogs are aggressively territorial (Pal et al. 1998;Bonanni et al. 2011;Pal 2015) should encourage further investigation of potential costs involved in this "migration" event and its repercussions. Monitoring the study dogs for a longer time post-event, especially in the mating and pup-rearing seasons (Sen Majumder and Bhadra 2015;Sen Majumder et al. 2016), might provide a clearer picture of the magnitude of the impact of such events on free-ranging dog populations. Nonetheless, the current study findings suggest a considerably prolonged significant residual impact of human overcrowding as a disruptive force on free-ranging dog dynamics. ...
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Interaction with its immediate environment determines the ecology of an organism. Short-lived perturbations in the habitat can adversely affect both wild and domesticated species. When such disturbances are unpredictable, they are more challenging to tackle. Therefore, specific strategies become essential for species to overcome adversities that can alter their dynamics. Species living in the proximity of humans experience significant anthropogenic disturbances. Response to such a disturbance, facilitated by human overcrowding, was studied in an urban-adapted species. Free-ranging dogs (Canis familiaris) found in developing countries can inhabit both urban and rural areas. Dogs present in urban habitats experience tremendous anthropogenic pressure. It is known that human flux can predict variability in their behavior in different ‘microhabitats’. We investigated the effect of a sudden massive surge in human footfall, during a major festival, on their abundance, distribution and behavior. While both abundance and general activity levels declined, vigilance behavior heightened during the festival. The behavioral activity was restored post-event, but dog abundance remained significantly reduced compared to pre-festival levels. Furthermore, we found variations in dog distribution across the sessions with higher dispersion during the festival. Conclusively, this study provides substantial quantitative evidence on the extent to which humans can influence even the behavior of animals well-adapted to urban environments. Significance statement Changes in the immediate environment drive organisms to build efficient adaptive strategies. These changes could be both natural, like flood and storm, or anthropogenic, like human overcrowding. The latter especially bestows immense stress on urban living animals, i.e., those that live in close proximity to human settlements. Using an observational study, we show that a sudden surge in human footfall negatively impacts a highly urban-adapted animal, the free-ranging dogs. Dogs lower behavioral activity, become more vigilant and modify their spatial distribution lowering their abundance in such areas. Though short-lived in nature, these disturbances can leave a long-term residual impact on their population. In conclusion, sudden human-induced perturbations can impact even those animals that are considered highly adapted to urban conditions. They are seen to adopt varied strategies to withstand such disturbances in their immediate environment, thus altering their behavioral dynamics.
... In some organisms, the timing of reproduction is mediated by lunar phase or tidal cycles (Fukushiro et al., 2011;Skov et al., 2005;Takemura, Sri Susilo, & SaydurMorita, 2004). In others, timing is strongly influenced by factors such as season, temperature and rainfall, which can result in a physiological state conducive for reproduction (Hirschfeld & R€ odel, 2011;Salminen & Hoikkala, 2013;Sen Majumder & Bhadra, 2015;Wikelski, Hau, & Wingfield, 2000). Biological rhythms, pheromones, visual cues and social facilitation can also play a role in coordinating behaviour (Gerlach, 2006;Kumar et al., 2010;Koizumi & Shimatani, 2016;Schiml, Mendoza, Saltzman, Lyons, & Mason, 1996). ...
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During the mass nesting of olive ridley sea turtles, Lepidochelys olivacea, thousands of adult females aggregate offshore, then emerge from the ocean together to lay their eggs along small, specific stretches of coastline. Little is known about the mechanisms that underlie synchronization of mass nesting. As a first step towards identifying environmental variables that control or affect the timing of mass-nesting events, we analysed 10 years of nesting data from Ostional, Costa Rica, a Pacific beach where mass-nesting events occur throughout the year. The onset of mass nesting was not uniformly distributed across lunar phase, with mass nesting occurring predominantly near last-quarter moon. Statistical models were developed to investigate whether environmental variables could be used to predict the timing and size of mass-nesting events. Predictions of the onset of nesting were improved by considering lunar phase and the time since the previous mass-nesting event as well as several oceanographic variables. Predictions of the number of turtles participating in nesting events were improved by considering salinity, relative humidity and nearshore current velocity, three parameters related to weather patterns and rainfall. Overall, the results imply that mass-nesting events at Ostional tend to occur near last-quarter moon but do not follow an invariant lunar or seasonal pattern. Instead, the results suggest that physiological and environmental factors interact to influence mass-nesting events, with no single environmental variable serving as an infallible predictor of either timing or size.
... Both in order to study free-ranging dogs and to compare their behavioural patterns with other canids, there is a need for an ethogram that is accessible to all. We have carried out detailed behavioural observations of free-ranging dogs in their natural habitats in India for several years and published partial ethograms focusing on certain aspects of their behaviour, like foraging, parental care, mating, etc. 4,19,[36][37][38] . ...
Full-text available
The domestic dog is known to have evolved from gray wolves, about 15,000 years ago. They majorly exist as free-ranging populations across the world. They are typically scavengers and well adapted to living among humans. Most canids living in and around urban habitats tend to avoid humans and show crepuscular activity peaks. In this study, we carried out a detailed population-level survey on free-ranging dogs in West Bengal, India, to understand the activity patterns of free-ranging dogs in relation to human activity. Using 5669 sightings of dogs, over a period of 1 year, covering the 24 hours of the day, we carried out an analysis of the time-activity budget of free-ranging dogs to conclude that they are generalists in their habit. They remain active when humans are active. Their activity levels are affected significantly by age class and time of the day. Multivariate analysis revealed the presence of certain behavioural clusters on the basis of time of the day and energy expenditure in the behaviours. In addition, we provide a detailed ethogram of free-ranging dogs. This, to our knowledge, is the first study of this kind, which might be used to further study the eco-ethology of these dogs.
Full-text available
Dogs, the most common companion animal of humans, perform not only the auxiliary of an individual, but also contribute to the nations’ crime and defence departments. Knowing the determinant-based disease status of dogs is imperative to keep them healthy by subsequent prevention and control of those diseases; however, such baseline epidemiological information is limited. Therefore, a retrospective cross-sectional study was conducted to estimate the proportional incidence of dog diseases; thus exploring the magnitudes in terms of intrinsic (ages, genders, breeds) and extrinsic determinant (season). Purposively, data of a total of 1557 cases of different diseases were collected from the record book of the Central Veterinary Hospital (CVH), Dhaka. Proportional incidence was calculated as proportion of cases of a specific disease among total number of cases of all types of diseases attended the hospital during the study period. Diseases of dog were categorized into infectious, non-infectious, and non-specific. Results showed that the highest proportional incidence was noted in infectious diseases (53.8%) followed by non-infectious diseases (23.4%) and the lowest in non-specific (22.7%) cases. Among them, disease-specific proportional incidence was remarkable in case of mange (9.5%), parvovirosis (8.7%), lacerated wound (8.5%), ectoparasitism (8.3%), helminthiasis (7.8%), and fracture (5.7%). The occurrence of mange varied significantly (p < 0.05) among all studied determinants (ages, genders, breeds, and seasons); while significant discrepancies (p < 0.05) in magnitudes of lacerated wound, dystocia, abortion, and gastroenteritis were observed among various age, gender, and breed, groups. Accordingly, dermatitis and orchitis had significant differences (p < 0.05) in proportional incidence amid various levels of age, gender, and season; whilst the burden of parvovirosis and alopecia differed significantly (p < 0.05) amongst different categories of age, breed, and season. The magnitude of otorrhoea showed a significant (p < 0.05) variation among different gender, breed, and season, groups. The proportional incidence of other diseases also varied significantly (p < 0.05) amongst either one or two studied determinants. This study provides a valuable insight about important diseases in dogs, which may serve as useful baseline information for disease prioritization and subsequent planning of effective control and prevention measures against those diseases.
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In 2018, Lindsay, Pluckrose, and Boghossian published four “hoax” articles within a number of disciplines that rely on critical theory (e.g., gender studies, feminism). When revealing the project, the authors argued that they wanted to expose these fields as being primarily motivated by ideology and social justice rather than knowledge generation. Their method tested the hypothesis that editors and reviewers will support papers that advocate “ludicrous” ideas including “fat bodybuilding.” In the pages of this journal, I presented a critique of their procedure, and the authors have provided a commentary on my article. After discussing the issue of whether their project was a hoax or not, I will argue that the crux of the matter is whether the papers were ludicrous/absurd. I will show how the authors made a fundamental error in their method; they failed to assess whether their ideas were indeed ludicrous/absurd.
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Co-habiting with humans in an urban ecological space requires adequate variation in a species’ behavioural repertoire. The eco-ethology of many urban species have been shown to be modified due to human activities leading to urban adaptations. Dogs (Canis lupus familiaris) are the first species to have been domesticated and have a long evolutionary history of co-habitation with humans. In the last two decades, scientists have investigated various questions on dogs pertaining to its domestication. In fact, no other species belonging to the family Canidae has received such attention in the scientific world. Unfortunately, majority of the work was confined to pet dogs in the western countries. Pet dogs are the result of artificial breeding based on desirable traits and their activities are typically determined by their owners. Free-ranging dogs found in most of the developing countries, on the other hand, represent a naturally breeding population without direct human supervision. Studying free-ranging dogs can thus provide us with crucial insights on the ecology and evolution of dogs in greater detail. Close to 80% of the world’s dog population is free-ranging, yet scientific studies on them are almost non-existent. Scientists have realised the importance and need of studying these dogs very recently to address various facets of the much debated domestication event. Free-ranging dogs are a highly successful urban-adapted species living in all possible human habitats in the developing countries. The dog-human relationship is highly complex and possess multiple trajectories. For example, these dogs depend on human subsidized food, choose dens near human households, yet receive a range of negative stimuli from humans; mortality of these dogs is mostly influenced by humans. In this thesis, we tried to answer questions relating to the dog-human relationship on Indian streets. My thesis involved an interdisciplinary approach where behavioural, cognitive, and ecological aspects are discussed to shed light on the evolution of the dog-human relationship. We began the work by looking at the natural history of free-ranging dogs in India. We collected data on the abundance of dogs and the distribution of their potential food resources, across India. Moreover, we recorded the sex ratio, group size, and behaviours of dogs at different study locations. We characterized study areas with regard to human activity levels by estimating human flux or movement and categorised them into low, intermediate and high flux zones. Our findings clearly suggested varying distribution of dogs and their food resources across different microhabitats in India. While a direct effect of food resource was not found, human flux significantly predicted the distribution of dogs. Moreover, we found a strong impact of changing human flux on the abundance and behavioural activity of free-ranging dogs. In the next section, we investigated the intra-group dynamics of dogs from the perspective of long-debated dominance-rank relationships. We looked at the steepness and linearity of agonistic and formal dominance hierarchies of groups of dogs from intermediate and high human flux zones. Our study did not reveal any clear dominance hierarchy among the free-ranging dogs, either in the intermediate or high human flux zones. The overall frequencies of interactions between the group members were found to be quite low, with many unknown interactions between for several dyads. We also proposed the use of subtle behavioural cues to maintain hierarchy rather than showing frequent behavioural exchanges in dogs. Findings from the study further led us to test free-ranging dogs’ interactions with humans. We found that these dogs interact with humans more compared to their conspecifics. Interestingly, we noticed that dogs rarely initiated behaviours towards humans, while humans played the predominant role in initiating both positive and negative behaviours towards dogs. We concluded that humans are a predominant part of the interaction network of the Indian free-ranging dogs. This opened up a window of testing dogs’ physical and social cognitive abilities. We found that free-ranging dogs lack the ability to persist on physical cognitive tasks and are poor performers like pet dogs. A higher dependence on humans is thought to be a key factor restricting dogs from persisting on an unfamiliar task. Interestingly, free-ranging dogs, as scavengers, showed competence while solving a familiar task, though task difficulty remained a factor that could not be disentangled. A partial dependence on humans was assumed to be the outcome of their long-history of co-evolution which resulted in a reduced problem-solving capacity in dogs. Surprisingly, a role of social facilitation was observed which predicted improved performances in both familiar and unfamiliar tasks. Free-ranging dogs like any other urban species are typically found to be aversive towards making direct physical contact with unfamiliar humans. The sociability of dogs was found to correlate with human flux, suggesting a role of life experience in shaping the personalities of these dogs. Dogs were shown to understand different human social cues and respond accordingly. The dogs in groups were bolder while responding to threatening cues from humans than in the solitary condition. Using two studies, we showed their ability to understand human pointing gestures, both simple and complex. The behavioural states of the dogs were heavily found to influence their responses towards humans. Dogs were found to be anxious or fearful while encountering an unfamiliar human. Interestingly, we found a crucial role of positive socialization in the form of petting in modifying such behavioural states of dogs and further building a strong dog-human relationship. In summary, this thesis provides unprecedented inputs into the current understanding of the evolution of dog-human relationship. The findings are not only restricted to the scientific advancement but may also be helpful in mitigating the growing doghuman conflict on the streets in India, by enhancing an understanding of the dynamics of the relationship between the two species, that might enable better management strategies.
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In this report, nighttime births of 3 stump-tailed macaques observed at the Aizawl Zoological Park, India, are described. Continuous focal observations were collected along with video and still photographs, on the 3 parturitions, from the first observed onset of labour. The average time taken for infant birth, beginning with visibility of the head at the vaginal opening, was 45 s. The births observed were similar in many re-spects, regardless of parity and social context. The average time taken for consuming the placenta was 4 min 4 s and the average number of contractions was 6.3. In all cases births occurred with the infant emerging in the occiput posterior position, assisted by the mother. Individual variations existed in the number of contractions, intercontraction intervals, self-examination of the anogenital region, duration of labour and the interval between infant birth and the delivery of the placenta. Each mother ingested the pla-centa completely, while holding her neonate, but without paying much attention to the neonate during placentophagia. Placentophagia appears to provide nutrition to the mothers. Detailed data on parturition in non-human primates, and particularly for Ma-caca arctoides, are still scarce. Data, such as those presented here, contribute to our un-derstanding of primate birth and the adaptive pressures that shape parturition behav-iour and reproductive success.
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Within the genus Loxia, it is among the crossbills that feed on the cones of pine trees (Pinus spp.) that the greatest morphological differences occur. Pine Crossbills also present the largest variations in the timing of the reproductive period. The presumed selective force in bill structure is considered to be cone morphology. Cones and scales of various species of pine were measured and a relationship sought with the bill characters of the species and subspecies of crossbills living on the various pines in the western Palearctic and in South-East Asia. Crossbills are renowned for breeding opportunistically and using the food supply in timing reproduction. We gathered data on pine cone predation during their annual cycle and on the phenology of crossbills breeding in relation with the maturation and opening of the cones. For the whole set of samples studied, no correlation was found between the variables describing the pine cones and those of the birds' bills. Principal component analysis (PCA) of the matrix of nine geographic locations * four morphometric cone variables and three morphometric crossbill variables led to a distinction between two subsets. A "cold" subset towards the negative pole included three bird species: Loxia pytyopsittacus and L. scotia (from northern Europe) and L. curvirostra from the Pyrenees, with a single pine species (Pinus sylvestris). A "hot" subset towards the positive pole covered Mediterranean and Asian locations; it involves one bird species; L. curvirostra and three pine species. So, at one end there are small cones and large birds (Scots Pines and northern European Crossbills) and at the other end there are large cones and small birds (Philippines). In these extreme situations of northern Europe and the Philippines, the structure of the bill does not therefore appear to be a character of adaptation to the size of the cones or the thickness of the scales. Parrot and Scottish Crossbills feed on the cones of the same Scots Pine as the Common Crossbill in the Pyrenees. The larger bill in the two former northern species could result from stronger selection pressure during the winter when the cones are closed and the seeds not yet ripe. On the other hand, in the Pyrenees, the cones ripen and open earlier making available large numbers of high-energy seeds in winter. Extra bill strength would not therefore be as advantageous as in Northern Europe. The Philippines Crossbill has the smallest bill size that is particularly noticeable on comparison with the Vietnamese subspecies, which uses the same Khasya Pine cones. This could be accounted for by the use of alternative food sources by the Philippines bird (mainly insects abundant in this tropical region) when the cones are closed and the seeds hard to reach. Statistical analysis of the central part of the PCA, corresponding to the Mediterranean, the Pyrenees and Vietnam (excluding the extreme locations: northern Europe and the Philippines) reveals a strong positive correlation between the proximal thickness of the scales and the index of curvature of the bill (r = 0.884; p = 0.019) in the six regions studied. The crossbills were all subspecies of L. curvirostra and the pines were of all the species studied. This supports the adaptive origin of the bill structure of the subspecies of Common Crossbills for the use of pine cones (excluding the insular form of the Philippines). The crossbills that live throughout the year in the pine forests can present two distinct breeding patterns: 1) strictly seasonal, limited to the period when ripe cones are available, as in northern Europe and the Philippines, 2) much more spread out, with an earlier start when the cones are still unripe as in southern Europe, North Africa and Vietnam. One of the originalities of the present work is to reveal a relationship between the structure of the bill and the breeding period. In crossbill populations where there is a correlation between the index of curvature of the bill and the thickness of the scales of the pine cone, the breeding period starts earlier and so lasts longer. This opportunistic capability of reproduction which becomes superimposed on the pattern of seasonality alone, presents numerous advantages for the population and, with the bill-scale correlation, indicates optimal adaptation to the use of resources in the different contexts: montane, Mediterranean and tropical. For crossbills in which the size of the beak is not correlated to the thickness of the cone scale, the length of the breeding season, and especially its earliness (before the cones are ripe) seems to be restricted by the availability of the food resources. Moreover the accessibility of food supply could have led to particular adaptations which can be morphological (large bill sizes in northern Europe) or behavioural (seeking alternative food supplies in the Philippines).
Chronicles the development of ion mobility spectrometry (IMS) from its inception two decades ago to current applications and plans for the future. Fundamental aspects of atmospheric pressure ion processes and their chemical ionization effect on the response characteristics of IMS are analyzed. The theory and physics governing the motion of ions drifting through an atmospheric pressure buffer gas in an electric field are presented in a simplified yet comprehensive manner. The book also presents an up-to-date report on recent developments and applications in IMS instrumentation, including laboratory instruments, monitoring systems, and hand-held instrumentation. The performance of GC/ IMS is analyzed and critically evaluated, and tables of reliable reducer mobility values are provided.
Reproduction in the pouched mouse (Saccostomus campestris) is inhibited in winter in the field and this seasonality is not controlled by photoperiod alone. The present study examines the hypothesis that reproduction in the pouched mouse is opportunistic (sensu Bronson 1989) and that the winter inhibition of reproduction is controlled by a reduction in either food quantity or quality. Food restriction (70 % of normal daily food intake) for six weeks, under short day conditions, was associated with a significant decrease in body fat index and mass of the accessory glands, testes and epididymides, while a high-fibre diet resulted in a significant reduction in fat index and masses of both the accessory glands and epididymides. A low-fibre/low-protein diet had no significant effect on either fat index or masses of the reproductive organs. Although the three experimental diets resulted in assimilation of significantly less energy than the controls, they did not inhibit spermatogenesis. We conclude that reproduction in the pouched mouse in the Eastern Cape Province is probably opportunistic and that the winter inhibition of reproduction may be controlled by a reduction in available energy which results from a combination of the lower ambient temperatures of winter and reduced food quantity and/or quality.