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Thank you for your thorough revisions. We are happy to recommend this manuscript for publication.
# PeerJ Staff Note - this decision was reviewed and approved by Nigel Andrew, a PeerJ Section Editor covering this Section #
Thank you for your careful revisions, which address the concerns raised by reviewers. The manuscript is now much clearer. Please address the additional comments provided by Reviewer 2 as well as these minor editorial suggestions:
Line 83 “Remove hyphen from “that-males”
Lines 87-89 Suggest replacing with, “…in species where the availability of mating partners can be very dynamic throughout the mating season, as is the case…” (i.e. move comma and remove “it”)
Lines 93-94 Suggest replacing “are still discussed until today” with “are still debated”
Line 189 Change colon to full stop
Lines 254-256 Suggest making this two sentences with the second reading, “We tested whether latency to begin mate searching and the probability….”
Line 315 – please add colon following subtitle to standardize formatting
Line 358 Suggest replacing “Except for this difference, Southern males also respond” with “In addition to this difference, Southern males respond”
Line 407 Suggest adding comma after “fitness”
Figure 1 Suggest replacing, “The tests were conducted in test areas (100 x 70 x 50 cm) in which males were exposed” with “Males were placed in test arenas (100 x 70 x 50 cm) and exposed”
Reviewer 1 requested that the conclusion be strengthened. This could still be improved, particularly the last sentence, which is rather vague (i.e. tie it to the current study).
Great improvement in the grammar and overall explanation of the manuscript.
The authors appropriately address the limits and constraints of their experimental design, while appropriately emphasizing the benefits of their design.
Data seems to be appropriately addressed, though some clarification regarding model comparisons (see below) would be helpful.
Line 54-56: When you list two or more items, you should make sure the list includes all of the same type of parts of speech (e.g., a list of nouns, a list of verbs, but NOT a mixture of nouns and verbs). Therefore, change to “or the consumption of time and energy”.
Lines 71-73: If they cannot perceive chemical cues, then how would they detect female density? This of course seems possible, but I suggest the authors suggest how this would be done (using a different sensory modality?)
Lines 263-264: “using the anova command” is a very R-specific way to describe what you did. Generalize what you did and describe the statistical test you conducted, not what command you entered into R. With model comparisons, using the anova command resulted in comparing the models using a likelihood ratio test. That the models were compared using LRT seems, to me, just as important.
Lines 264-266: Perhaps I am misunderstanding what you are stating here. As currently phrased, if you were to retain “the model with the highest p-value”, this makes it sound like the simplified model and the original model each were given a p-value after the anova command. This is, of course, not true, as the LRT results in a single p-value. Of course, if p<0.05, the simplified and original models are different, and one of them is a significant improvement upon explaining variation in the data. I would focus on the latter (that you chose to keep a model if it successfully explained more variation in the data (unless you used AIC comparisons instead, which needs to be noted if the case).
I am also not sure what you mean when you say that you stopped when you “only found significant differences between the models”. Wouldn’t you stop when there were no significant differences between the models?
Line 280: Please change to “The sample size of surviving males was fairly small, which was why we only conducted a Fisher exact test…”
Lines 368-369: What makes you think this would be especially true if the information is chemical-based?
The topic of this paper is interesting and covers a gap in knowledge. In particular, both reviewers and I appreciate your candid discussion of your data and analyses. Both reviewers have provided useful feedback and have made suggestions for improvements. Additionally, below are a number of mostly editorial suggestions to improve the readability of your manuscript. Please try to address all of these comments and amend the manuscript as appropriate. We look forward to reading your revised submission.
Throughout the manuscript, please refer to “virgin females” as “adult, virgin females” or “sexually mature, virgin females” because all females used here are virgins.
Line 46 Change “missing” to “miss”
Line 59 Change “in mating systems, where” to “in mating systems where”
Line 70 Remove “so far”.
Line 73 Replace “Different” with “Other”
Line 91 Remove comma in “mating system, in which”
Line 92 Consider changing “wrong” to “poor”?
Line 111 Change “as they are” to “because they are”
Line 112 Change “and a reduced” to “and reduced”
Line 133 Change “around” to “approximately”
Lines 134-135 It appears the PeerJ preference is “sp.”, not “spec.” and not abbreviated.
Line 135 Consider rewording to “Spiders were provisioned with water six…”
Line 135 Change “On this occasion” to “on these occasions”
Line 143 Change “moulted” to “moult”
Line 145 Change “design) and offered” to “design), which offered”
Lines 148-149 Suggest changing to “We used the tibia-patella length as an index for body size.”
Line 150 Remove hyphen in “measuring-tools”
Line 173 Suggest merging this section with the section above “Experimental design”. “Experimental conduction” sounds awkward and this really is more information about how the experiments were designed.
Line 199 Suggest changing “for the data analysis” to “for data analyses”
Lines 221-222 I am struggling with the logic of this test. What is this actually testing? If you wanted to test whether the male would be more attracted to the largest female, shouldn’t you have place the male equidistantly to all females?
Line 238 Please add stats after “(Table 1” so that the reader can compare easily to the stats given earlier in this sentence.
Line 262 “Unlike predicted neither” needs editing. Suggest “Contrary to our predictions, neither…” or something along these lines.
Line 269 “Different scenarios”… shouldn’t this be about alternative explanations for the same scenario? Please reword. Can you expand this paragraph to discuss these alternative explanations more fully?
Line 274 Suggest changing “claim” to “suggest” or something similar
Lines 274-287 This paragraph describes a reasonable argument why female density should be important, and more so in Northern Europe than in the Mediterranean. This left me wondering why samples from Southern France were used in your experiments. Perhaps it would be useful to explain why they were chosen here?
Line 288 “We argue” instead of “we argued”?
Line 289 Suggest adding a comma after “availability”
Line 301 Please clarify “males rather perceive virgin then immature or mated females” – do you mean they more easily perceive them?
Line 321 Change “unknown, whether” to “unknown whether”
Line 350 Remove “at least”
Line 356 Change “The diversity” to “This diversity”
Please add your statistics in parentheses in Figure legends.
#25218
Mate availability does not influence mating strategies in males of the sexually cannibalistic spider Argiope bruennichi
Author(s): Anna-Lena Cory & Jutta M. Schneider
MAJOR ISSUES
(1) 1 experiment with 2 treatments or 2 separate experiments?
(2) Use of the term “[word(s)] availability”.
(3) Experimental Conduction (LINES 173-196): Experimental setup and running of the trials.
- - - - -
LINE 73: “The causes of sexual cannibalism are discussed until today and seem to be diverse.”
> Citation needed. Suggestion = (Wilder et al. 2009).
Wilder SM, Rypstra AL, Elgar MA. 2009. The importance of ecological and phylogenetic conditions for the occurrence and frequency of sexual cannibalism. Annu. Rev. Ecol. Evol. Syst. 40, 21–39.
LINE 112: “…extreme female-biased sexual size dimorphism…”
> How do you define “extreme” female-biased sexual size dimorphism? Suggestion = see (Schwartz et al. 2013, Hormiga et al. 2000, Scharff & Coddington 1997).
Schwartz SK, Wagner WE, Hebets EA. 2013. Spontaneous male death and monogyny in the dark fishing spider. Biology Letters, 9, 20130113.
Hormiga G, Scharff N, Coddington JA. 2000. The phylogenetic basis of sexual size dimorphism in orb-weaving spiders (Araneae, Orbiculariae). Syst. Biol. 49, 435–462.
Scharff N, Coddington JA. 1997. A phylogenetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae). Zool. J. Linn. Soc. 120, 355–434.
THROUGHOUT THE MANUSCRIPT
Be consistent with word/term use.
Specifically: low-availability vs. low mate availability, high-availability vs. high mate availability
LINES 173-196: Experimental Conduction
The specifics of the experimental setup (“types” of arenas) and running of the trials (starting point, front end, back end) were difficult to follow.
25218 - RAW DATA
Column B: density = high-availability, low-availability?
Column F: male_adultweight = units?
Column G: male_testweight = units?
Column M: female_adultweight = units?
Column N: female_testweight = units?
Column T: copulation_duration = units?
Why is V20 (test_no) 1425 (male_id) highlighted (Column M & X)?
As the manuscript is currently written, it is not clear if the research is covering: 1 experiment with 2 treatments (high-availability & low-availability) OR 2 separate experiments: Experiment 1 = High-Availability (4 virgin females), Experiment 2 = Low-Availability (1 virgin & 3 sub-adult females).
If the authors choose to go with 2 separate experiments, I would suggest they split their MATERIALS & METHODS and RESULTS into “Experiment 1” and “Experiment 2” with appropriate titles/headings.
It is important to publish negative results, however I think the current manuscript would be more publishable (i.e. stronger) if an additional experiment (or two) and/or relevant natural history observations were added.
LINE 14: “While many studies focused on male competition, the impact of female density has been widely neglected.”
> While many studies HAVE focused on…
> The impact of female density on what?
LINE 18: “Males mutilate their paired genitalia to prevent sperm competition…”
> To REDUCE sperm competition? Do we know that it completely prevents it?
LINE 20: “Males can reduce the risk of cannibalism if they jump off the female in time, but will then transfer less sperm.”
> FEWER sperm, since fewer is used with words denoting countable things (e.g. sperm).
LINE 21: “An alternative solution of THIS trade-off is to copulate longer, commit self-sacrifice and secure higher minimal paternity. THIS self-sacrificial strategy may be adaptive if prospective mating chances are uncertain.”
> Use THIS.
LINE 31: “We found that males needed more time to start mate searching if the mate availability was low.”
AND
LINE 37: “However, we found no evidence that different levels of female cues and mate availability…”
> Both PERCEIVED mate availability?
LINE 33: Remove “often”.
LINE 37: “However, we found no evidence that different levels of female cues and mate availability prompted adjustments to mating decisions in males.”
> Suggestion: However, we found no evidence that different levels of female cues and availability prompted adjustments to male mating decisions.
LINE 39: ONE reason…
LINE 52: “…in response to a high risk of male competition…”
> Consider using STRONG male competition as used later in LINE 110.
LINE 52: “…mate guarding, nuptial gifts, or ejaculates…”
> Consider using AND/OR, since can’t it be a combination?
LINE 54: “…the effects of female density often remain unnoticed…”
> Unnoticed by who?
LINE 67: “Here, it was found that males developed faster if they received volatile…”
> Here? Where? Are you referring to a specific species or study system?
LINE 71: “In many mantises and spiders, males show an extremely high limitation in mating rates…”
> Extremely high limitation in mating rates? Do you have any examples of this?
LINE 73: “Different studies HAVE found that sexual cannibalism depends…”
LINE 87: “…also function as mating-plug…”
> Mating-plugs.
LINE 97: “Females stereotypically attack males during copulation, but males can reduce the risk of cannibalization if they copulate short and try to jump off the female in time.”
> Suggestion: Females stereotypically attack males during copulation, but males can reduce the risk of cannibalization if they copulate quickly and try to jump off prior to female attack.
LINES 117, 146, 148, etc.
> Are these “mating” or “choice” tests?
LINE 133: “On two days a week…”
> Two days a week…
LINE 143: “Females of both maturation states were put in little frames…”
> You list the dimensions. Remove “little”.
LINE 157: “In both treatments, males were exposed to four females, but we manipulated the number of receptive females.”
> Receptive = Pheromone Emitting females?
LINE 165: “…we used individuals from Southern Europe…”
> Specifically France?
LINE 174: “In the time of July 14th until August 16th, 2016, the experiments…”
> Suggestion: Between July 14 and August 16 2016, the experiments…
LINE 175: “…underneath a waterproof pavilion roof.”
> So was this an open-air space?
LINES 176, 181, etc.
> Are these “choice” or “test” arenas?
LINES 175-177: “The tests were conducted in cleaned choice arenas that had a size of 100 x 70 x 50 cm. The floor was covered with packing paper that was exchanged after each test.”
> Suggestion: The tests were conducted in clean arenas (100 x 70 x 50 cm) with the floor covered with fresh packing paper.
LINES 199, 200
> We used 3 times. Consider revising the use of “We used…”
LINE 208: “…which we log-transformed the copulation duration.”
> Log-transformed why?
LINE 225: “In the low-availability treatment, males left the starting point significantly later...”
> Left the starting point = started mate searching?
LINE 227: “…reintroduced to the starting point after 45 minutes…”
> Why?
LINE 244: “The only significant predictor of sexual cannibalism was male size (Table 2, Fig. 2).”
> What was the predictor? Was it larger or smaller males? Specifically mention it in this sentence.
LINES 360-366: Conclusion
> The CONCLUSION seems a bit weak. Consider revising it.
LINE 361: “Male mating effort is often traded off against future reproductive success and should strongly rely on prospective mating opportunities.”
> Consider revising. Something like… Male mating effort is often a trade-off between A and B.
FIGURE 1 CAPTION
> Effects of WHAT on copulation duration? Add more detail to the figure caption so it is able to stand-alone.
FIGURE 2
> X-axis label? Males?
TABLE 1 CAPTION
> Testing the effects of WHAT on copulation duration?
TABLE 2 CAPTION
> Testing the effects of WHAT on cannibalism rate?
Overall, the authors' organization of their paper is to be applauded. The figures, tables, and literature references appeared to be well-thought out and appropriate. There were many cases, however, where the grammar needed to be dramatically improved.
Some examples of grammar that need to be fixed include the lines below. However, I eventually made most of these comments in the General Comments section instead. Eventually, I admittedly got a bit tired of fixing some errors, so I politely ask the authors to take the changes I have suggested and be sure to apply it to all areas of the manuscript, and not just the lines mentioned. I also want to point out that there were, in fact, many paragraphs and sentences that were quite well-written.
In the ‘Background’ section:
Line 14: Please change to “While many studies have focused on male competition” (add “have”).
Line 15: I suggest having the 3rd sentence use the same verb tenses. “Here, we aimed to close this gap and tested effects of female availability…” (change aim to aimed)
Line 23: Please change to “We hypothesized that males opt” (change hypothesize to hypothesized). First, it should arguably be in the past tense, because you already made the hypothesis before conducting the study. Second, you used the past tense in the last sentence of this section (“we further predicted”), so it should, at the least, remain consistent.
In the Results summary (at the beginning):
The word choice could use some improvement.
Line 31: I politely suggest changing “needed more time to start” to something like “males took more time to start”.
Line 32: I would also re-structure the 2nd sentence to avoid stating “low mate availability led to”. Instead, state that copulation duration and sexual cannibalism rates were not affected by manipulations in mate availability.
In the Discussion summary (at the beginning):
Line 36: You state that the availability of virgin females facilitates mate searching, but not other mating decisions. Please re-phrase this, as many would consider mate searching to be a type of mating decision. Alternatively, re-phrase “mating decisions in males” to something more specific.
The experimental design appeared to be entirely original and appropriate for the hypotheses that the authors proposed. There were certainly some shortcomings that I pointed out in my General Comments, but in many cases, the authors already appropriately addressed these shortcomings in their Discussion.
I do, however, strongly suggest that the authors make several of the changes that I have proposed in the General Comments to improve the overall framework of their study, given the experimental design of their study.
The authors do a good job of providing alternative approaches and explanations as to why they might not have found significant results. In fact, I strongly believe that this paper is exactly why it is important to still make sure that non-significant data gets published and is available to the scientific literature.
Some data analyses comments are made in the General Comments, along with some possible additional explanations or thoughts to consider regarding why you found mostly non-significant differences.
Overall, the authors did not overly speculate. Some changes, however, should be made regarding more clarification that they are studying current context, not the effects of past social experience.
This is the section that has the majority of my comments and suggestions in it. Please take the time to address each of these thoughts, suggestions, or requests. Respectfully, many of them do need to be improved before, in my opinion, the manuscript should be considered for acceptance.
Introduction:
Overall, the first paragraph could use some improvement. The authors do a good job introducing the first sentence. However, the rest of the first paragraph needs citations and additional background information. Otherwise, it almost reads more like a prediction (especially the last sentence of the first paragraph).
Line 51: Is there a review paper that you can cite to back up your statement that most studies focus on the latter? If not, perhaps consider changing the sentence to “although many studies focus on the latter”.
Second paragraph: This paragraph is well-written and includes useful, informative citations and background information. Good job. I would consider ending it with a more conclusive “take-home” message though. Perhaps something like “Therefore, that might be some scenarios in which female density is the only social cue that can be accurately assessed.”
Line 64: Change “All three taxa” to “Some members of these taxa are known”. Otherwise, the reader might think that EVERY single member of the taxa can use chemical cues to assess female quality. Additionally, I would consider using a different word than “quality”. Perhaps, use traits, condition, or status. Too many sexual selection biologists dislike the word “quality”.
Lines 65-66: Please make this a new sentence. No need to follow up a statement about all 3 taxa, then switch to discussing only spiders.
Line 69: Perhaps, more specifically, you mean “whether past chemical information about female availability is also used…”
Line 72: It looks like you forgot to include the 2nd author on the Elgar citation. Please fix.
Line 73: Please fix the verb tense. I think you mean “the causes of sexual cannibalism are still discussed today and are diverse”
Lines 85-86: The phrasing makes it sound like all spider species only have two maximal mating chances. This is, of course, not true, so please fix the phrasing. The same is true for the next sentence in Lines 86-88.
Line 97: Please change “if they copulate short” to “if they copulate briefly”
Lines 100-110: Citations are needed for several of these facts and statements about the study species. Please add them.
Lines 115-124: Good job. In terms of the information previously provided in your Introduction, your predictions make perfect sense. I also appreciate the careful use of words, as you are correct, you are not testing “female density”. You are testing “mate availability”. On a side note, this is comparable to studies that have investigated sex ratios or operational sex ratios – the operational part includes only the mature, reproductive individuals, while sex ratios include all individuals, regardless of their reproductive status.
Materials and Methods:
Lines 129-130: A quick look at Cory & Schneider (2018) suggests that very little details about the lab breeding strategies are described. Specifically, can you please provide the reader with details regarding whether or not the spiders used in your current study are siblings or not? How many different eggs sacs are represented by the spiders in your current study? This is important information that will inform the reader about how transferable and repeatable your results might be.
Line 133: Please remove “On” from the beginning of the sentence.
Lines 134-135: Please look into PeerJ’s editing preferences. I am not certain whether “spec.” is an appropriate abbreviation for species. I am much more familiar with “sp.” (singular) or “spp.” (plural). Either way, in one instance you have it italicized, while it is not italicized in another.
Line 135: I am more familiar with providing water to spiders (and other arthropods) ad libitum through the use of cotton wicks or rolls. Can you please provide more detail about how you provided the spiders with water? Did you simply spray their web? Why 6 times a week and not 7? I am simply curious.
Lines 136-141: As an arachnologist, I am familiar with these terms and the differences between penultimate vs adult male and female genitalia. Many readers, however, will not. Could you please provide some citations that might direct the readers to the original (or more recent) description of the genitalia for your specific species? If such a description does not exist, at least a more general description for orb-weaver spiders or Argiope as a genus would suffice.
Lines 158-159: Make sure it is clear that you are talking about your species only. Recently, a study was found that demonstrated that widow males could copulate with penultimate females and successfully reproduce using this alternative mating tactic.
Lines 163: Please explain how ages were matched. Does this mean some batches of 4 virgin females had an average age of 21 days post-maturity (but were all “matched”), while others had an average age of 10 days post-maturity (but were all “matched”)? Matching does not seem as important to me as making sure that the average age across groups was not statistically different and was randomized.
Lines 166-168: Please provide more details about what is actually being measured here. Specifically, when you state “males copulated with similar large females in both”, it sounds like copulation (no or yes) was the dependent variable. Please re-phrase so it is more clear as to why you needed to run a t-test. (I am confident I understand why, but the phrasing can be improved to better make your point).
Line 168: You are still assuming, though. Therefore, please change the word from “conclude” to “assume” or something less strict.
Lines 169-171: Respectfully, this is a bad habit to start doing, though many behavioral ecologists, unfortunately, do so. To the best of my understanding, you did not statistically incorporate (which is a very difficult thing to do in the first place, because not all females were re-used) the fact that some females were re-used, while others were not. I applaud the authors for being honest. There is a possibility, however, that the re-used females did not make contact with the males in previous trials for a reason. You might be unintentionally biasing the results by continuing to use individuals that produce negative results (in which there could be an array of reasons why).
Lines 186-189: On the same day, they were re-tested? Why not test them on a different day? Further, many would argue that not making a choice, is in fact, a choice. Are you not unintentionally forcing them to make a decision if you give them a 2nd chance any time there is not a decision that is made?
Lines 195-196: I politely, but strongly, disagree with the authors’ operational definition of bigyny here. Why is a male considered bigynous if he left, but simply did not return? Should not bigyny be only used to describe males that actually mate twice? Perhaps you need to re-adjust your definitions. Alternatively, I might politely suggest to change the phrasing to “bigynous mating strategy”. That way, it suggests that they might be bigynous, but you technically do not know for sure. One option will require you to re-do your statistical tests, but from my perspective, some sort of change is necessary.
Lines 207-208: I highly suggest using a survival curve, so that you do not have to log-transform your data. I am not entirely sure if you had a cut-off point (e.g., if they were copulating for more than a set amount of time, you had to end the trial), but if so, this would also allow you to “censor” your data to statistically incorporate that.
Lines 209-210: Please use a citation in your description of body condition. There are a variety of ways to define this, and taking the residuals is one of these ways. Please add an appropriate citation (e.g., Jakob et al 1996).
Results:
Line 221: “Closest female” was never described to be incorporated into your statistical models in your Methods section. Please provide the details as to how you tested this in your Methods.
Lines 228: Your p-value is greater than 0.05, therefore, this sentence is not accurate.
Lines 234-238: I appreciate your disclosure in how you addressed the outlier. It would be tempting to ignore it, or never mention it in the first place. I applaud you for simply presenting both stats results and allowing the reader to decide how to interpret that result. Good job.
Lines 248-250: Again, please re-phrase your definition of bigynous. On a related note, do we know how long it might take a male before he would mate a 2nd time with a new female?
Lines 262-268: I was delaying this observation to wait and see if the authors eventually addressed this. I suggest that the authors strongly emphasize that they are studying decision-making within the current context, not based on past social experience. For example, the Bailey et al. 2010 paper that is cited in the Introduction provided experience with male courtship signals to test the effects. From the best of my understanding, you did not provide experience with different availabilities of potential mates. You manipulated the current context. I highly suggest digging into the literature deeper regarding this difference and to think about (and ultimately, discuss in your paper) whether or not that is why you did not find any significant results for many of your variables. Perhaps they need more experience with these social cues in order to demonstrate plasticity in their mating behaviors. Specifically, look at Kasumovic & Brooks 2010 and read about socially-cued anticipatory plasticity. Also, take a look at one of my own papers, Stoffer & Uetz 2015, entitled “The effects of social experience with varying male availability on female mate preferences in a wolf spider”. I have a series of other manuscripts that have also focused on plasticity in mate preferences in Schizocosa ocreata that would perhaps be of interest as well. While I am not trying to make a shameless plug for my own research, I think this paper could prove to be rather useful for your paper to consider and discuss. It focuses on the female decision-making end, however, and not the male decision-making end. It still, however, should be a good read on examining this difference between past experience and current context.
Lines 278-281: Studies on Schizocosa (Hebets 2003, Rutledge et al. 2010, Stoffer & Uetz 2015, 2016, etc.) might also be good to take a look at regarding asynchrony of maturation.
Lines 290-291: While I understand why you chose your method, I still think it would have been useful to also include a treatment group in which there was only 1 adult virgin.
Lines 294-298: Given that this was the case, do you know what % of your subadults were in fact penultimate females? This is a very important component of your study and could explain why you did not obtain significant differences! (Though I understand your follow-up paragraph) In short, females could prefer virgin females when given a choice, but it could still affect their mating tactics in a different way.
Lines 312-319: Great point. Nice paragraph.
Lines 344-346: Again, when you state sentences like this “the effect of mate availability on male investment”, I immediately assumed they were given experience with different treatments of mate availability and subsequently measured for their mate investments/tactics. I think the phrasing throughout the manuscript needs to make this difference a bit more clear.
Lines 353: If my memory serves me correctly, the Berglund 1995 paper is another great example of a study that technically measure the current context of manipulating female density, not the experience of it. In reality, these organisms would be experiencing these social manipulations, not have zero experience and then suddenly be thrusted into a low-availability or high-availability context. Your study still is very interesting and is necessary to our understanding. I am just giving you other perspectives to think about and address.
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