Behaviour17 Conference

Scientific conferences are some of the biggest highlights of my year. I just attended the Behavior2017 Conference in the beautiful seaside town of Estoril, Portugal.

Pictures Colour Library  - PCL

Estoril: warm sun, cool breeze and seawater

I’m still early enough in my scientific career that when I attend a conference, I often meet, for the first time, people whose work I’ve read. Public speaking is always a bit scary, but it’s such a joy to come together with people from all around the world who are working on similar, related questions in different organisms. There is not enough time to go over all the talks, but here’s a small sample.

Jonathan Pruitt gave an engaging talk on something he calls “social susceptibility” which is the extent to which the collective behavior of a social group is affected by a “leader” or keystone individual. In his remarkable social spider system, there are different discrete behavioral types (personalities), and introduction of a single extremely bold aggressive individual can make the entire colony more aggressive (more individuals attack). But this effect only occurs in deserts, not in the less dry savannah, because the aggressiveness of a colony is only advantageous in deserts. Pruitt tested whether the social influence of the bold individual (the leader) is determined by the traits of the leader or the followers. And apparently, it’s the followers that matter, not the bold leader. In deserts, the shy followers are influenced by a bold individual from either a desert or a savannah or even a heterospecific spider. In savannahs where aggressiveness is not adaptive, the shy individuals were not influenced by this extreme individual. Pruitt ended by making a general (obviously political) metaphor that influential “leaders” can only have be influential if the surrounding society as whole follows them, and varying levels of this “social susceptibility” can exist from group to group due to different selection pressures. As Pruitt put it, given high enough social susceptibility, a society can even be greatly influenced by a “heterospecific sham” (I found that bit hilarious). It’s a fun and complex story, and Pruitt told it well.

Another favorite talk was by my friend Neeltje Boogert on the effects of early life stress on social learning in zebra finches. Using a series of clever experiments, she and others found that when solving foraging “puzzles” young birds copy their parents, but when the young were given stress hormones, they stopped copying their parents and instead copied unrelated adults! Neeltje then did a similar study with song learning and found that young males tended to copy the song of their father, but the experimentally stressed juveniles did a worse job copying their father’s song. The stressed males had weaker father-son bonds and they may also have been more likely to copy non-father males. I wonder if similar phenomenon exist in humans: do highly stressed kids start learning earlier from their peers rather than their parents?

Another great talk was by Amiyaal Ilany, also about parent-offspring relationships in the context of social networks. Ilany had previously worked on a model exploring the idea of maternal inheritance of social networks (something I hope to test in vampire bats when I get a large enough sample size). Here, Ilany applied this idea to a huge long-term dataset collected by the Holekamp Lab from wild hyaenas. He showed that a hyena does indeed have similar network ties to her mother. This similarity depends on maternal social rank, in that offspring of mothers of higher rank inherit a stronger similarity to their offspring’s network position. The social inheritance of bonds also depends on the amount of time that the individual spends with their mother. One of the great thing about this study is that it shows where social network structure comes from.

I finally met Thomas Bugnyar (even if only for a moment). He gave a terrific talk on social cognition in corvids. He showed that ravens adjust their calling behaviour according to who was in the audience. The victims of an attack from another raven called more when their kin and bonding partners were present but called less when friends of the aggressors were in the audience. I’m a big fan of his work on social cognition and sociality in ravens and the corvids themselves.

Rene van Dijk gave a great talk on vocal kin signatures and kin discrimination in social weaverbirds. I’m hoping to look at paternal vs maternal kinship signatures in calls soon in vampire bats; I’ve been collecting data on this since 2009.

Sue Healy gave a great talk on the how birds build nests. There’s surprisingly many decisions involved about materials, colors, shapes and structures, and again, how early life experience can affect nest-building. I always love reading or hearing about cognition experiments, because the experimental designs are always so clever.

Barbara Taborsky discussed evidence from their cooperatively breeding cichlids about how a subordinate uses either helping or a submissive display to prevent aggression and eviction by dominants. The submissive display might somehow honestly signal that there is no challenge to the dominant. Interestingly, although the strategy can vary based on the immediate social context, there is consistent individual variation in strategy use. But it is not heritable. Instead, it comes about through something like a lifetime specialization into either ‘helper’ or a ‘submissive’ type based on early life experience. This later influences whether the subordinates either disperses, or stays and helps at the nest.

There were many other studies that manipulated the early environment and looked at effects on social behavior. For example, in zebrafish early environments influence social competence and gene expression in the brain years later.

Michael Taborsky gave some interesting new results on different cooperative strategies employed by cooperatively breeding cichlids, including reciprocity. I’m excited to see those papers when they come out. I’m hoping to do some experiments with reciprocity in rats next year.

There were several terrific posters from University of Exeter on cooperativeness in predator-inspecting guppies. They seem to have a really fantastic department there for animal behavior.

I enjoyed Dustin Rubenstein‘s “big picture” talk about comparing animal societies (including the often ignored taxa like spiders, thrips, and shrimps) to understand social evolution with some neat results from his work on snapping shrimp. It was great to finally see what these snapping shrimp look like! I’m really impressed with the number of systems that his lab works on in the field.

I had an interesting discussion with Dieter Lukas and Corina Logan about brain size evolution (see their recent review manuscript) and the necessity for open access (and other changes to publishing in science).

Yvonne Zurcher had an interesting poster showing evidence that vocal similarity influences the development of pair bonds in common marmosets (in addition to call convergence once bonds form).

Frans de Waal gave an entertaining talk with fun videos of nonhuman primates cooperating.

I have better appreciation for the idea of indirect genetic effects after this meeting.

Hanna Kokko, a leading theoretical biologist in evolution and ecology, gave a talk that changed the way I thought about evolutionary bet-hedging (which I mentioned at the start of my talk on “social bet-hedging”). I was a bit worried she would not like my rather loose use of the term “bet-hedging” but instead she seemed quite interested in my idea of how individuals invest in social relationships, which was very exciting for me. When we spoke I was a bit too nervous and star-struck actually. I think I stammered something like “I really love your book on modeling. But I’m really sorry…. I did not finish it…. I only made it to Chapter 3 when I was in grad school…” [And then maybe I trailed off because my brain was screaming: WHY ARE YOU SAYING THIS?]. Alas, this is one of those embarrassing moments that I’ll think about later randomly and shudder.

Barbara Koenig described her recent work on sickness behavior and its effects on social networks. This is of great interest to me, because we are now doing similar experiments in the vampire bats, led by Sebastian Stockmaier. The first paper should be out next year.

I took notes on far more talks and posters, but this blogpost could go on forever, so I’ll stop here.

There were several more prolific people I wanted to meet such Redouan Bshary and Frans de Waal. But so often I think to introduce myself, they are talking with someone else, and I started to think to myself, Do you actually have anything to say other than ‘I love your work!’ ?”. Then I would think, “What’s a good question I have for him?” … and then I start overthinking and then the moment is gone, hahaha. If I read enough papers that I like by someone, they eventually (and quite irrationally) begin to seem like some sort of celebrity to me. This is strange because I don’t generally like the idea of celebrities, focusing on people rather than ideas, but… it still happens. For better or worse, academia is just inherently social, with so many of phenomena that are being discussed at this very conference:

  • dominance hierarchies
  • networks of association with individuals greatly varying in centrality
  • leaders and followers
  • early career experiences that set individuals on different paths and specializations
  • reciprocity and other strategies for making cooperative investments while avoiding exploitation,
  • and (of course) “social bet-hedging“– the strategic tradeoff between investing in strengthening of existing relationships and creating new ones


Conferences give me a boost of inspiration. One of the main reasons I love conferences is that I get a lot of ideas while I’m sitting there listening to talks, or from the talk itself, or from conversations with people. In my crazy busy life at Panama, I almost never have time to just sit and think about ideas; I’m always caught up in the details of a particular experiment.

The next meetings I hope to attend are something called “CBEN 2017” and this one, both on social complexity and cooperation.



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molecular evolution in vampire bats (video)

More information: Sears Lab


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English translation of Park (1988) Dominance relationships in a colony of vampire bats. Korean Journal of Zoology.

Rachel Crisp is currently writing her masters thesis on dominance interactions in female vampire bats. Males have a clear dominance hierarchy in competition over roost territories, but do the female vampires have a dominance rank? If so, does it strongly correlate with cooperative interactions? Her work should come out next year!

Despite all the work on cooperation, not much is known or published about competitive interactions in female vampires. Interestingly, female vampire bats are larger than males, but there is no evidence for a strong female reproductive skew as in cooperative breeding societies. The little bit on competitive interactions that is available is in German (work in the 1970s supervised by Uwe Schmidt) or Korean (by Shi-Ryoung Park). Rachel Moon graciously translated this 1988 paper by Park from Korean to English.

For easy accessibility, here it is below.

Dominance relationships in a colony of vampire bat, Desmodus rotundus

Park, S.R.

Korea National Univ. of Education, Chongwon (Korea R.). Dept. of Biology

Translated by Rachel Moon, Harvard University

Originally published as Park, S. R. (1988). Dominance relationship in a colony of vampire bat, Desmodus rotundus. The Korean Journal of Zoology. 31 (4): 243-250.



Dominance relationship was investigated in a captive of Desmodus rotundus, a neotropical sangivorous bat, under seminaturalistic conditions. The hierarchy was determined from four different behaviors (flee, fly-out, avoid, wait) by the encounter of two adult bats on the feeding site. The aggressive action “flee after fighting” was relatively low (16%) compared to the other three observed behaviors. A hierarchy of the females was reflected sometimes in the feeding order. The harem male dominated the non-harem males and exhibited his territorial behavior. However, to his female partners he didn’t show aggression.



Except for a few species, most bat species (Chiroptera) live socially (Gopalakrishna, 1955; Eisentraut, 1957; Kulzer, 1958; Goodwin and Greenhall, 1961; Barbour and Davis, 1969). Bats exhibit complex social structures and diverse kinds of social behavior (Bradbury, 1977). However, the functional significance of bat social structure and social behavior is still not well studied; therefore, this study aims to understand the significance of complex social behavior of bats by studying dominance relationship in a group of South American neotropical vampire bats (Desmodus rotundus).

Vampire bats (Desmodus rotundus) have developed and adopted unique behaviors and physiological structures that enable them to feed on the blood of vertebrate animals. Vampire bats form long-term small colonies with highly developed social structures (Wimsatt, 1969; Schmidt et a., 1978; Wilkinson, 1985). Until now it has been nearly impossible to study dominance relationship structure of vampire bats in natural settings; therefore this study aims to investigate dominance relationship in a group of vampire bats in experimental settings that resemble naturalistic conditions.



This study was conducted with 9 vampire bats (Desmodus rotundus; three males, six females). Only one male and one female were captured in the wild (Colombia, 1975), and others were born and raised in captivity. The bats were kept in a thermostatic chamber (with 12h/12h of light and dark) at a temperature of 25C and 70% humidity. They were provided cow or pig blood (with removed fibrin) on a bird water feeder. In order to distinguish each individual from another, I marked each bat by attaching aluminum loop of different colors on its forearm (Table 1).

The observation cage had a size of 250 x 180 x 100 cm, and its left side, right side, back side and ceiling were made with thin plastic surface with small holes so that the bats could hang easily. The front center part of the room was made with transparent glass (100 * 50 cm) so that the experimenter can observe easily.

In order to observe the bats’ behaviors, an infrared night vision scope (metascope 9902E) was used, during early hours (17:30-20:30) of the 12 dark hours. This study emphasized an understanding the quantitative aspects of behavior observed in a previous study of vampire bat social relationship (Schmidt and Manske, 1973). In other words, I recorded and analyzed frequency and duration of observed behaviors, distribution of behaviors, and subsequent behaviors.

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Table 1


The social behavior of bats takes place at a feeding site, where they often encounter one another. During the initial period of feeding time, aggressive physical fights were often observed. This behavior usually begins with two bats pushing and shoving each other. If the opponent does not back off immediately, this aggressive physical fight occasionally turns into a fierce combat. To first determine the feeding dominance relationship of bats in the colony, we investigated the feeding times of individual bats, as seen in Figure 1.

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Figure 1

D (female) always feeds first; this female kept her place until she was full, even though other younger bats who were feeding at the same time often obstructed her by physical fight of pushing and shoving aside (Figure 2A). H (female) always showed up at the feeding site after the first feeding activity period, when other bats were mostly not feeding. During this period, H often encountered her 15-month-old daughter J at the feeding site. J would often push H to the side, although J had several unoccupied feeders nearby. Then J would wait right behind her mother H until H finishes feeding and returns (Figure 2B). As soon as H left the feeding site, J started to feed at the same spot H was feeding.

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Figure 2 (sadly unclear) and Table 2

However, not all of the other colony members ate their meals following such straightforward feeding order. Depending on each feeding site situations (whether bats are at the feeding site or not, and if so which bats are present), the bats visited the feeding site at irregular intervals. Therefore, it was difficult to determine group dominance relationship solely based on feeding order. Because the bats frequently encountered different individuals at the feeding site, it was possible to observe several types of social behavior. I chose four different types of social behavior exhibited by two encountering bats as the main parameter to determine bats’ dominance hierarchy.

  • Flee (after fight): After two adult bats meet and fight at feeding site, the winner bat continues feeding or starts to feed, whereas the loser bat flees away. For fighting behaviors, I observed shoving and pushing to the side, and attacking behavior.
  • Fly-out: It is a behavior shown when the inferior bat immediately flies away without showing any aggressive behavior when two bats encounter at the feeding site.
  • Avoid: When an inferior bat is feeding and another bat approaches, the inferior bat gives up its meal and moves to a different feeding site. The inferior bat feeds at a new feeding site or stands for awhile at a distance.
  • Wait: The inferior bat waits at a nearby place, or goes back and forth hesitantly at a different place, until the superior bat finishes feeding and leaves.

Fleeing is exhibited after aggressive behavior, whereas fly-out, avoid, and wait behaviors are “direct-flight” behaviors performed by inferior bats themselves in order to avoid potential conflicts; therefore, these three types of behavior are characterized as unaggressive or submissive behavior.

A total of 294 encounters were observed at the feeding site, and the frequency of each behavior type is shown in Table 2. Aggressive behavior accounted for 15.6% of all observed behaviors, which shows that it happened relatively less frequently than unaggressive behavior (Figure 3). Aggressive behaviors between male bats at the feeding site rarely took place; the most common behavior between two encountering male bats was “fly-out” behavior, which accounted for half of all behaviors exhibited.

In order to determine dominance relationship, I used a point system in which the superior (winner) bat earns 1 point when superior bat and inferior (loser) bat are distinguished. The score difference between two bats is shown in Table 3 by comparing the scores of each bat. Using this result to build a sociogram of dominance hierarchy at the feeding site, the following phenomena in each group could be explained.

Dominance relationship among harem females (Figure 4)

When encounters took place between two females from five harem females, the most common social behaviors were “wait” and “avoid.” D was the most superior dominant bat among five harem females. B, who had the lowest dominance hierarchy ranking, exhibited “wait” and “avoid” behavior as a response to D’s behaviors. E had a middle dominance hierarchy ranking, which was higher than inferior bats C and B and lower than superior bats D and H. E responded by “fly-out” behavior all three times when she encountered H. Although I never observed direct encounters of H and D at a feeding site (their feeding period almost never overlapped), I concluded that D had a higher dominance hierarchy ranking than H depending on feeding ranking.

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Table 3, Figure 3, and Figure 4


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Figure 5

Dominance relationship among males (Figure 5)

When two out of three male bats (G, A, F) encounter at the feeding site, the inferior bat usually flees away. Thus a clear dominance hierarchy was revealed among these three males. The most superior bat was G; whenever A or F appeared at the feeding site, G forced them to leave the site. When G approached, the inferior bats rushed their feeding. The most inferior bat was F; whenever F was feeding, I often observed him looking around his surroundings and rushing his feeding. Similarly, A also exhibited typical feeding behavior of inferior bats, which was carefully approaching feeding sites, quickly feeding non-stop, and leaving in a hurry.

Out of 31 encounters between G and A at the feeding site, A showed “fly-out” response 26 times; of 42 encounters between G and F, F responded by flying out 32 times. Out of 12 encounters between A and F, F exhibited “fly-out” behavior in all encounters. Therefore it was determined that the most superior bat was G, the most inferior bat was F, and A was in the middle.

Dominance relationship among male bats and harem females (Figure 6)

At the feeding site, G (male) always showed passive behavior towards old harem females who have previously given birth to pups. When these females were feeding at the feeding site, G responded by exhibiting “wait” behavior 30-40 cm away from the site until the females finished feeding. I observed six encounters between G and B (female who hasn’t yet given birth), and four out of six times B showed submissive behavior (“fly-out”) towards G.

A (male) showed dominance towards B and C (females); however, the number of encounter between A and D or H (females) was too small to clearly determine their dominance hierarchy.

F (male) was certainly superior than B and C (females), who had low dominance hierarchy rankings within harem females. However, he ended up getting an inferior status after frequent pecking-order disputes with other harem females. His dominance hierarchy ranking was determined by his fights (“fleeing”) with other females, which were 19 times (73%) out of 26 encounters.


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Figure 6


Vampire bats (Desmodus rotundus) form small groups when they go out to feed in the wild. Greenhall et al. (1969) observed behaviors such as combats, waiting, and co-feeding when vampire bats were feeding in groups in the wild. Furthermore, even in captivity the vampire bats often engage in pecking order disputes at feeding sites; therefore, the social structure based on dominance hierarchy seems to be a prominent characteristic of social behavior of vampire bats (Schmidt and Van de Flierdt, 1973). Vampire bats maintain close relationships through various social interactions at feeding sites, such as combats and submissive behaviors. According to the results of this study on a captive group of vampire bats, aggressive behavior (16%) took place less frequently than unaggressive behavior (submissive behavior), and most of the bats showed defensive or submissive behaviors (wait, fly-out, avoid) at the feeding site.

Park (1986) reported that aggressive behaviors are usually triggered by subadult bats, showing a strong tendency to claim high dominance status. Park observed that bats who are 14-16 months old often engaged more in dominance ranking fights compared to adult bats. The results of this study also revealed that group members abide by dominance hierarchy through a single consistent order when feeding and that aggressive behaviors between adult bats at feeding sites did not occur as often. Thus, it can be thought that vampire bats have especially well-developed mechanism of group formation, which serves to restrain mutual aggression by strengthening the cohesiveness among group members.

Vampire bats form a typical harem social structure (Load, 1976). Such social structure has been observed in other bat species such as Pipistrellus nanus (O’Shea, 1980), Phylolostomus hastatus (MaCreacken and Bradbury, 1981), Carollia perspicillata (Porter, 1979), and Artibeus jamaicensis (Morrison, 1978); the male bats in these species also form a single distinct dominance hierarchy, which is observed by ritualized combats. Park (1986) found out that alpha male (harem male) vampire bats stayed with females for a long period of time, whereas beta males stayed at a place where females use as their temporary shelter. On the contrary, gamma males stayed far away from female groups and did not exhibit any territorial behavior. Connecting to the results of this study, it seems that hierarchy system of male vampire bats is associated with territoriality. In the context of dominance behavior, Wicker and Uhrig (1969) observed Lavia frons engaging in a territorial fight with neighboring bats, and Bradbury and Emmons (1974) witnessed Saccopteryx leptura driving out outsider bats from its hunting territory. Several other bat species (Vespertilionidae: Dwyer, 1970; Brosset, 1976; Phyllostomatidae: Fenton and Kunz, 1977; Porter, 1979) are known to defend their harems and exhibit territorial behavior towards intruders and competitors.

Sex and age of bats are significant factors in formation of dominance hierarchy. Usually, older animals show dominance over younger animals, and males show dominance over females (Immelmann, 1983). This study reveals that alpha males (harem males) did not exhibit absolute dominance over females; on the other hand, harem males exhibited inferior behaviors toward their female partners. It is not easy to declare dominance relationship based on age only with the results of this study. Whether juvenile bats depend on dominance hierarchy rankings of their mother bats require further investigation.


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Updates (July 2017)

New timeline

  • July 30- August 4, 2017: I’ll be at the Behavior 2017 Meeting in Portugal
  • September 3, 2017: Team Vampire wraps up captive experiments and we begin our fieldwork led by Dr. Simon Ripperger.
  • October 15, 2017: I leave Panama.
  • October 18-21, 2017: I will be at the North American Bat Meeting (prospective students who are attending–> email me)
  • November 1, 2017: I move to the Max Planck Institute of Ornithology in Germany
  • August 15, 2018: I move to Ohio State University, USA and start our new lab!!

Recent and relevant papers

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Some advice for grad students

With my new lab starting in Fall 2018, I am now interested in prospective graduate students and postdocs. More information on how to apply here. Below are some of my thoughts and advice on applying to graduate schools and being a graduate student.

Check out this advice written by wiser people than me:


On choosing a graduate program

Rather than basing your decision on the prestige of the school, choose a good lab (research group led by a PI). The best schools tend to have better labs but not always, and the most prestigious schools might have a more competitive atmosphere, which may or may not be your thing. Schools can specialize by hiring several excellent faculty in a specific area, such as animal behavior. For example, University of Texas Austin and University of California Davis have two of the best animal behavior programs in the world, despite not being as famous as Harvard or Oxford. The name of the school does matters and so do the other labs in your department, but for a research career in our field, what matters most is the quality of your papers not the prestige of the school. So the question should be: which lab will help me do my best possible work?

A good place to see a bunch of labs at once is to go to a scientific conference. Don’t be afraid to ask a senior grad student or postdoc: If you were me, what lab would you join?

Don’t apply to the school until you’ve talked with the PI. Also, talk to at least three people in the lab and ask them what the PI is like. Are they hands-off or demanding? What are lab meetings like? How quickly do they respond to emails? Ask them in a way and in a setting where they can be totally honest. Expect to hear different things, but look for common themes.

The PI matters a lot, but so do the other lab members. You’ll probably spend more time with them than the PI. So try and evaluate about the culture of the lab. If the PI has a very large lab, see if there’s a postdoc that could be your direct mentor.


On doing a masters before a PhD

In some countries and at some schools, getting a masters before a doctorate is mandatory, and I think there’s actually some sense in this. But many undergraduates in the USA who want to do a PhD in biology think getting a masters in biology does not make sense and is a waste of time. I’d like to present the counter-argument. There are two good reasons to get a masters before a PhD.

First, doing a masters puts you one step ahead when you start your PhD. Becoming a good scientist typically takes a long time. Many students think that a masters degree puts you two years “behind” but it actually puts you two years ahead, because a masters is like a “practice PhD”. You’ll get better at critical reading, writing, and statistical analysis. If all goes well, you publish a paper, maybe even two. When you start your PhD, you’ll be a better researcher than you would have been coming straight from your undergrad. A 2-year MSc degree can even put you on track to get your PhD two years earlier, or get you a faculty interviews as postdoc two years earlier.

One secret to success in academia is trying to be one step ahead of where you need to be at each stage of your career. As an undergrad, that means publishing a paper. As a starting PhD student, it means having a compelling proposal to apply for fellowships. As a finishing PhD student, it means establishing yourself as an international expert on the topic of your dissertation. As a postdoc, it means laying the foundations for a fundable long-term research program. At each stage, you will not be compared to others based on your age. So while impressive, there’s no real long-term advantage of being a 24-year-old with a PhD. Instead, you’ll be competing for funds with others at the same career stage, based on what you’ve done so far. My point is that what you know, and what you’ve done, are more important than how quickly you get various degrees.

Second, doing a masters gives you time to decide if academia is right for you. Being a research scientist in a field like behavioral ecology is one of the most creative and intellectually stimulating jobs one can imagine. If I had a million dollars, almost nothing about my work life would change. But science is truly a “labor of love”, and academia is neither the fastest nor easiest path to securing a permanent, stable job. For instance, 12% of college baseball players go on to play in the major leagues, whereas only 10% of incoming biology PhD students become biology professors. As an undergrad, you can get a pretty rosy picture of academia talking with professors. There’s survivorship bias: the people who tell you about academia as a career are the few people who succeeded in it through some combination of talent, perseverance, and luck. Professors are not a random sample of grad students; whereas many grad students suffer through graduate school, many professors look back at their grad school years with sweet nostalgic memories of having the time to focus solely on their research. It’s easy for professors (especially older professors) to think, “I simply did X, Y, and Z and that’s how I became a professor”, but today, the statistics for becoming a professor are actually a bit grim. Each year in the USA, there are 16,000 new biology PhD students. Of those successful recruits, 63% get their PhD, and the average time to that degree is 7 years. Of those graduates, 70% get a postdoc and of those postdocs, 30% get a second one. Only 15% of postdocs get a tenure-track job within 6 years (~13 years after receiving a PhD). Most starting PhD students in biology in the USA want to be a tenure-track professor. But only about 7% of them actually get that job.

If you’re unsure about doing a PhD, just ask yourself this question: if you knew you had a zero chance of getting a faculty job, would you still want to spend 7 years doing a PhD just for the experience itself? If the answer (as it would be for many of us) is: “Yes! I love doing research“, then of course do a PhD! You cannot lose. If you’re unsure, then perhaps consider a masters instead. Then, you’ll be in a better position to know whether to commit to the longer academic journey, complete with greater expectations, fewer deadlines to structure your time, and an extra dose of imposter syndrome. Some starting grad students realize that they don’t want to be in academia in the middle of their PhD, but it’s difficult psychologically to quit and do something else that makes you happier. That transition can feel like failure. In contrast, you can always turn a MSc into a PhD, and if you start with the goal of getting a masters, you can leave with a masters and that is a great success. It’s a way to test the waters.


On writing research proposals

Brainstorm a list of topics and questions that are truly interesting to you. Study the most fascinating topic you can. Read the literature, but don’t let the literature tell you what questions are interesting. That is, don’t get brainwashed by other people’s research agendas: it’s good to think outside the box.

Start with a compelling question, then think about how someone could get the answer, then look to see if anyone has done exactly that.

Many big interesting questions are addressed by one or more general theories and every theory makes key assumptions and predictions. Have these actually been tested yet? That’s a good place to start. For example, biological market theory assumes that a shift in supply or demand will lead to changes in the value of services exchanged by individuals in a mutualism. Has this been tested? Reciprocity theory predicts that individuals will reduce cooperative investments towards individuals that don’t reciprocate. What’s the evidence for that?

Not every interesting phenomenon is directly addressed by a scientific theory, but often there is a theory, perhaps from another discipline, that could be applied to the problem. That’s another good starting place. For example, what theories from biology could be explain the psychology of human friendship? If your research doesn’t address a theory, it should address a big question that is of interest to many people.

Your proposal should start with the big picture. This should be something that everyone will find interesting. Answer these questions:

  • What’s been done before? Show that you know the background work. Show that you know what studies are relevant and which aren’t.
  • What’s the problem with the current state of knowledge or the current paradigm and the evidence for it? What’s missing? Show you can think critically.
  • What needs to be done to fix that problem? What could be done by you in just the next few years? Show that you can set realistic goals.
  • If [hypothesis], then [prediction]. Show that you can think logically and clearly.
  • How exactly will you test this prediction? Don’t be abstract and vague. Be concrete. Paint a picture. Get the reader to think: “Ok, I can definitely see this happening”
  • What resources do you need to accomplish this?
  • If applying somewhere: Why is location X a good place to get those resources? Why is person X the best person to help you? Demonstrate that you know the person or lab.
  • Why can we be sure that you will finish the task?

After you write the draft, get lots of feedback. Get input from as many minds as possible. The most likely scenario is that inexperienced researchers will propose to do ambitious things X, Y, and Z in their first year, and more experienced people will tell them to start off by focusing just on part 1 of X. Just doing that might take 2 years.


On managing your PI

Scientists don’t receive any training in how to be a good mentor. Your PI will be committed to your success but they won’t necessarily know what you want or need unless you tell them. Nobody wants to be “needy” but it’s a good idea to be proactive. If the PI doesn’t check up on you, send them monthly updates. When you write outlines and drafts, ask for feedback. If you want to discuss things, ask them when is good to meet. Professors have packed schedules, so it’s usually better to set a meeting, then to just drop in and expect to have a long discussion.


On being a good scientist

Being a successful academic is not always the same thing as being a good scientist. There are academics who are terrible scientists, and there are great scientists who are not very successful in academia. Being a successful academic means being a good teacher, getting large grants, and publishing in prestigious journals. In sum, it means that you have lots of influence in your field. Being a successful scientist, however, means that your influence is actually moving your field towards a more accurate view of the world, because your work is careful and rigorous, you encourage other people to critique it, and you are honest about the limits of your conclusions. In sum, you say things that are actually true. Science is a job for people who value intellectual honesty, skepticism, logic, and evidence. Science is inherently open and transparent. If you try to succeed in academia at the expense of the quality of your science by overselling your work or making straw man arguments, you may do well in the short-term, but you will eventually gain a bad reputation as a sloppy scientist among the leading researchers in your field. And those are the people you should care the most about impressing.

Ideally, we are both good scientists and good academics. The ideal lab has a culture that encourages being a good scientist first and foremost, by trying to create an environment where everyone feels safe to be ignorant and ask really naïve questions; where undergraduates feel comfortable arguing with the PIs, postdocs, and grad students; and where nobody criticizes people, but we all feel comfortable criticizing ideas (constructively). Not taking criticism of ideas personally or defensively is one of the one most difficult, yet important, skills for scientists. We must remember that the goal is to make the final product as good as possible, not to be the most expert or clever person in the room. Likewise, our goal should be to produce work that is transparently valuable, not to “get it past the reviewers”. Even if criticism is not constructive or just plain wrong, it still tells you what parts of your argument you may need to communicate more clearly. Expertise only comes from failing, a lot.

The whole point of academia is meeting and talking with talented, interesting, and passionate people who know a lot about something, so do not be afraid to seek out, talk to, and listen to more experienced people who can think more clearly than you about topics with which you’re struggling. Yes, more senior scientists always seem too busy, but there is nothing they love more than to use their arcane knowledge to really help out a scientist in training. Looking back, I realize I wasted so many opportunities to talk science with really smart and knowledgeable people. Yes it can be embarrassing to reveal just how little you really understand about a topic, but everyone expects that from a student, so now is the best time!










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Team Vampire, Spring 2017

Katharina Eggert is from Germany and visited STRI from March until May 2017. She helped with a broad variety of projects including scoring cooperation in vampire bats, maintaining a system of monitoring bat roosts, and measuring exploration of novel objects by young and old vampire bats.


I have always had an interest in the ecology and conservation of the flora and fauna of the tropical rainforests. Before interning at the Smithsonian Tropical Research Institute with Dr. Gerald Carter, I had worked at field stations in Ecuador and Peru. My interest in bats started in spring 1994 when a small colony of Pipistrellus pipistrellus moved into a crevice above my window, and decided to return every year thereafter. I have spent many summers watching “my” bats and they motivated me to join and volunteer for our local conservation organisation. During my internship at STRI I had a unique opportunity to study cooperative behaviour and social bonds in the common vampire bat. I learned about designing and evaluating research projects that focus on social interactions. In addition, I assisted with a range of different bat-related projects in Gamboa and Barro Colorado Island. This enabled me to gain a wide range of valuable bat-relevant field skills.



Hugo Narizano is from France and visited STRI from February to May. He is currently a MSc student at Edinburgh Napier University with Jason Gilchrist, and is studying the relationships between self-grooming and social grooming in vampire bats.

My interest in biology, especially ethology and wildlife, started as a child when I was living on a former farm in a small village. Growing up surrounded by a wide range of different animals, including some mysterious bats, triggered my desire to devote my life to animals. I started to be particularly interested in the similarities between humans and non-human animal sociality. This project was my first opportunity to work with bats, and with the common vampire bat, which I consider to be one of the most interesting bat species. I am learning more about designing and running experiments. Also, this project has enabled me to meet intelligent and inspiring people from STRI studying different topics and species. I have learned more about science as a whole. This project has taught me about what it takes to become a competent scientist.

After I graduate from Edinburgh Napier University, I am willing to conduct another research project with vampire bats, or on a different topic with bats, such as the interaction between Nepenthes hemsleyana, a pitcher plant, and Kerivoula hardwickii, the Hardwicke’s woolly bats [see past blogpost]. In the near future, I also wish to follow on with a PhD focusing on bat behaviour or ecology preferably, or possibly on a eusocial species.



Jineth Berrío-Martínez (MSc in Biology) is a researcher from Colombia who arrived in April has been travelling around Panama to find additional wild vampire bat colonies. She is working on development of biting ability in young vampire bats. 

My research interests include tropical biology, population ecology, reproductive biology, evolution, and conservation. Early in my career as a biologist, bats caught my attention, leading me to work with them for more than eight years now. Desmodus rotundus is one of my favorite bat species because of the astonishing morphological and behavioral adaptations related to their diet. Despite the fact that vampires are a common species in my home country, there are many aspects of their ecology that are unknown or poorly understood.

Without a doubt, this internship has been an extraordinary opportunity to gain experience in behavioral ecology. Gerry is teaching me about experimental design and statistical analysis. Additionally, I hope to provide valuable support to a variety of different projects from filming and measuring behavior to fieldwork. I intend to make sure that my research projects are completed with diligence.

Many of the most remarkable scientific papers I have read on tropical ecology, behavior, and evolution have been written by researchers associated with Smithsonian Tropical Research Institute (STRI). Here I am exposed to other researchers’ experiences studying ecological and evolutionary problems. I strongly believe the skills I will learn and the collaborations I will establish during this internship will be an important platform for achieving my future academic goals. I plan to improve my research skills and pursue a PhD in Ecology and Conservation abroad.




Samuel Kaiser (MSc in Biology) arrived in June from Germany and is currently conducting a thermal preference experiment in vampire bats. The goal is to see if captive-born vampire bats without any past experience have an innate preference for warm blood.

It is hard to pinpoint my interests. I like animal behaviour – how animals communicate, interact and socialise. I’m also interested in how animals are affected by the environment and humans. Previously, I studied microplastic pollution and its effect on Daphnia magna. Then, starting an internship at the Max-Planck-Institute for Ornithology (Seewiesen, Germany), I discovered my fascination for bats and their use of echolocation to orient and hunt. After conducting experiments on passive-listening in the greater horseshoe bat (Rhinolophus ferrumequinum), I switched my focus to studying thermoperception in the tropical bat Phyllostomus discolor. This ultimately lead me to the internship at Smithsonian Tropical Research Institute in Gamboa, Panama with Dr. Gerald Carter and Dr. Rachel Page to learn more about the common vampire and its preferences for warm blood. My fascination with the common vampire is based in its ability to sense infrared radiation. This internship is a great opportunity for me to study tropical bats, and I have always wanted to do fieldwork in the tropics. In the future, I plan to travel around Central America and later start a PhD in animal behaviour.


Sebastian Stockmaier is a PhD student at the University of Texas at Austin in Dan Bolnick`s lab. This is Basti’s fifth field season in Panama and his second season working on vampire bats. He is studying the influence of sickness on cooperative behaviors in vampire bats.  basti

Despite my passion for being outside and for all the wildlife out there, I chose to spend most of my undergraduate research in laboratories working on virus-cell interactions (Institute Pasteur in Paris) and phylogenetic classification of some mysterious African bat viruses (Robert Koch Institute in Berlin). I then started a Masters program in organismal and evolutionary biology at the University of Konstanz, where I worked with Dina Dechmann and Teague O`Mara on several bat and bird species, as well as on tiny European shrews.

My masters at the University of Konstanz taught me–besides the fundamentals of evolution, ecology, and behavior–about the beauty of getting out into the field. Some formative influences included several research projects at STRI in Panama, and a spontaneous research trip to Kasanka National Park in Zambia, a spot known among bat researchers for a massive seasonal gathering of flying foxes. Having experienced biology in both the field and lab, I am trying to bridge the gap in pursuing some of my research questions. For my doctoral degree, I joined Dan Bolnick’s lab at the University of Texas. One of the main questions in our lab revolves around how pathogens interact with their hosts , a topic which involves ecology, behavior, and evolutionary genetics. I am generally interested in how transmissible pathogens shape group-living, and how they influence the evolution of social traits. I am also interested in how social behaviors of hosts can shape the evolutionary trajectories of a pathogen. I am trying to tackle some questions using vampire bats, and for other questions, I plan to establish experiments in Austin using aggregating C. elegans genotypes and a nematode pathogen. Since I enjoy teaching as well as research, I would like to further pursue a career in academia.


Along with Basti, Rachel Crisp has also worked in both 2016 and 2017. She is studying social dominance in female vampire bats.

“Team Vampire” is part of the Rachel Page Bat Lab at STRI.


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Should we wipe out vampire bats?

Adam Cole from NPR visited our lab to shoot this great video short on human-vampire bat conflict.

So “should we wipe out vampire bats?” No, even if we could, we shouldn’t try, and frankly, nobody is actually suggesting exterminating vampire bats as a long-term solution. More on this topic in a previous post:

Is killing vampire bats good for public health? No. It might increase rabies risks.

The best work on this topic is being done now by Daniel Streicker’s lab.

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