What's the point of authority if you can't use it to crush dreams?

I've just been looking at extant species (so ignoring extinct things and hybrids) in the comparison. So as a hybrid this falls outside of the domain I've been curating

This taxon is being used confusingly for identifications. Observations are being identified as it both for situations where it is an definitely an observation of a unisexual hybrid individual (treating it as a hybrid taxon), and if it's impossible to separate from pure individuals (treating it as a complex taxon). In my area, where it is impossible to separate pure Jefferson's Salamanders and unisexual individuals, half of the observations are ID'd as this taxon and half are as Jefferson's. If this taxon is actually meant just for the hybrids, then my interpretation is that technically all of our observations should be at genus level. However, the taxon name "Hybrid Ambystoma 'unisexual complex'" is ambiguous and lends itself to being used for both purposes.

It that inconsistency fine? Other options would be just having a complex taxon that includes all the species involved, or having that and a species/hybrid level taxon. The biology itself is confusing so I don't know... https://www.inaturalist.org/observations/7923194

weird, I just got a notification that said:
"deleted user resolved a flag for Ambystoma 'unisexual complex' 7:56 AM" (central time)

met too - not sure what that was all about

I'm in favor of swapping Ambystoma jeffersonianum into Hybrid Ambystoma 'unisexual complex' if others are

@loarie I will let @pantherophis comment further, but I do not believe this is a good proposed change.

2/2

Since most of the challenges involve Blue-spotted/Jefferson and LLJ/LJJ unisexuals, I would suggest making the following changes.

-“Blue-spotted Salamander” should have subtext added that says “or A. laterale-dependent unisexual”.
-“Jefferson Salamander” should have subtext added that says “or A. jeffersonianum-dependent unisexual”.
-Retain the unisexual taxon as “Indeterminate Ambystoma sp. unisexual”. There are sometimes intermediate or ambiguous specimens which are clearly not one of the “pure” species, and while there would be no doubt that they are a unisexual, it may not be possible to say with certainty to which complex they belong. This is more accurate than defaulting them to “Ambystoma sp.”

This will handle the majority of the unisexual-related ID issues that crop up. The justification for this is because most of the time, while you can’t be sure if it’s unisexual or not, the unisexual indicates the presence of the pure species. At least in Ontario, the convention among field biologists is to simply refer to a specimen as “Blue-spotted” or “Jefferson”. The existence of the unisexual complexes is understood, and the qualifier “or a [species it looks like]-dependent unisexual” is implied. I think this would convert this convention into an iNaturalist format, so to speak, and allow a more orderly and more useful sorting of salamander observations.

Known exceptions could simply be noted in observation fields by those who are familiar with the specific situation. There is at least one LJJ population in Ontario that is actually dependent on A. laterale. That situation may or may not be anthropogenic. However, it would be easy enough to ID those as “A. jeffersonianum or Jeff-dependent unisexual” and note in a field that this is from that rule-breaking population. This is still important to separate – in this case, Ontario ascribes equal legal protection to both pure Jeffersons and LJJs as species at risk. Identifying either – or anything that appears to be either – may convey automatic habitat protection, and if it isn’t an already known population, it would warrant further investigation by scientists able to test the DNA.

I suspect it would not be desirable to add the “or [species in question]-dependent unisexual” tag to A. tigrinum, since over most of its range I don’t think this is an issue. Again, observation fields could be added for known situations. It may be an open question for A. texanum. Certainly there are many areas where it only occurs as the pure species, but where they do occur, unisexual lineages involving A. texanum can be messy. On Pelee Island in western Lake Erie, there are relationships between A. laterale and A. texanum that may be unique. Among other things, “LT” diploid unisexuals are known that resemble A. texanum but that are dependent on A. laterale. If it isn’t too much trouble, I would suggest a special taxon for this, a “Pelee Island A. laterale-A. texanum unsexual complex”. This should take care of the majority of A. texanum-related issues that are posted here.

The pure species here are real entities, and the various unisexual lineages are real entities, and there needs to be some way to acknowledge them within the limitations of the platform. If this, or something like this, could be done, it would help organize the data into useful categories with a reasonable degree of certainty and accuracy. Quick example of how this can help: the actual, current distribution of the Jefferson Salamander in Ontario is poorly established. Its “official” status has fluctuated over the decades - mapped as widespread, then described as absent. The current "horseshoe" map is relatively recent. It is quite likely that populations exist that have yet to be documented. This is something for which iNaturalist is perfectly suited. If phenotypically Jeffersonian individuals are identified as such, while they may in fact be LJJ unisexuals, it still identifies areas with probable Jefferson populations, which in turn helps to guide further investigation and conservation efforts. If anything Jefferson-ish is dumped into a generic unisexual taxon, important discoveries of conservation significance may not come to light.

While I think this is long enough, I’m able to expand on the various points in here if needed.

I think not enough relevant people are aware of this flag conversation, but having more people involved would confuse things as well. There are so many ways of dealing with the situation. Should we tag more identifiers of these?

Does putting all the species involved in a complex taxon and just identifying ambiguous individuals to complex level oversimplify the situation?

Is it only Blue-spotted/Jefferson that have most of the problems? They're the only 2 in the group we have here so I'm ignorant of the others.
If that's the case, then we wouldn't need a complex for all of the species, just those 2. Or a complex within a complex, not sure if that's possible or not...

@pantherophis there isn't any function to add subtexts to species names. Do you mean like just adding that comment to the end of the default common name? Not sure how tolerated that would be.

There is no consensus yet that I can see. @pantherophis seems to basically be suggesting leaving it as it is but getting people to identify differently.

The problem is that people are using the taxon as both a complex and as a taxon. It actually represents like 5(?) "taxa". pantherophis is suggesting that identifiers identify by phenotype rather than than genetics. So if it looks like a Jefferson's Salamander, call it a Jefferson's Salamander even if it could be a unisexual. Only identify clearly intermediate unisexual individuals as the unisexual taxon.
He also suggested adding another “Pelee Island A. laterale-A. texanum unsexual complex” but I'm not sure why it would be helpful to have this in addition to the “Indeterminate Ambystoma sp. unisexual”?

I think it would be helpful to eventually make more subsets of the complex. In Illinois we've got 'Silvery' salamanders which a lot of people count as its own species as they have been breeding amongst themselves for many decades, so the population is becoming more distant from the rest of them. Same thing up near Chicago, the Unisexual moles have been breeding with blue-spotteds for decades and they are more different from other populations. I've heard people refer to them as the 'Chicago Mole Salamander'. I may be reading this wrong but that's my two cents.

@brdnrdr sorry didn't see your response until mine was finished.

In Illinois we've got 'Silvery' salamanders which a lot of people count as its own species as they have been breeding amongst themselves for many decades, so the population is becoming more distant from the rest of them.

My understanding is that these salamanders can't breed parthenogenetically, so they need a population of males of the pure species. Which species is involved here, and can they be distinguished from the unisexuals? (I'm guessing Jefferson's if it's "Silvery"?) Same thing with the Chicago "Tremblay's" population.

In southern Ontario there are multiple populations where people have no idea if they're seeing Jefferson's or "Silvery", so they end up getting identified as either/both.

I guess you could distinguish the Tremblay's from Blue-spotted by size. Presumably there are pure female Blue-spotted there as well though?

First, and most importantly, it isn’t accurate. There isn’t one type of unisexual. There are at least four lineages, maybe more
...
Of the four species involved here (Ambystoma laterale, A. jeffersonianum, A. texanum, A. tigrinum), each one can be reliably identified from one another when distinguishing among pure species.
...
Retain the unisexual taxon as “Indeterminate Ambystoma sp. unisexual”. There are sometimes intermediate or ambiguous specimens which are clearly not one of the “pure” species, and while there would be no doubt that they are a unisexual, it may not be possible to say with certainty to which complex they belong. This is more accurate than defaulting them to “Ambystoma sp.”

@pantherophis given these statements from your earlier comments, would there be any issue with just eliminating the ambiguous "unisexual complex hybrid" taxon and merging it into an "A. laterale complex" taxon containing all 4/5 pure species? Any unclear unisexuals could just be identified to complex level.

We could create a new taxon (should it be species or hybrid?) for the Pelee Island unisexuals if they are distinctive enough (or just leave it since people could always filter for complex-level observations from Pelee Island).

You all seem to be more knowledgeable about unisexual Ambystoma than I am. I think I’ll be good with however you all decide to handle this.

Fair enough, thanks both for your input.

To add my two cents, from a "layman's" perspective:

Initially, I was IDing almost any Ambystoma sp. as a unisexual, unless it was clearly a male (since males would 99.9% be the "pure" species).

In speaking with others, I concurred that functionally, this was not helpful - even though potentially "more" correct. I have since then been sort of "policing" the identification using these guidelines:

Obvious males would be ID'd as the pure species (in my case, this is always A. jeffersonianum or A. laterale due to my location).

Strong indicators of a diploid/unisexual such as limb gigantism, would be ID'd as a unisexual. This is still not 100% verifiable in my opinion.

Anything else would be ID'd as the pure species (again, in my cause this is almost always a Jefferson's or Blue-Spotted)

I add my comments here from a practical sense. I can't add anything more, and certainly am not in any position to be able to dispute, information presented here by others. What I can say is that making the Genus easy to identify, and to keep it as the most helpful to any researchers, is probably the best approach.

I believe this following statement is a fact: There is no way to identify a "pure" species from a "unisexual" species from images of larvae, juveniles or adults salamanders with 100% accuracy. The only way to truly identify these is through lab work, or I believe in conjunction with looking at egg masses in the case of Blue-Spotted and unisexual (I believe the former does not lay noticeable egg masses, so egg masses would indicate a unisexual).

From my perspective, the complexity of having so many different species to identify would not be practical. I would propose lumping them together, at least for the 2 core species we see most often:

a) Blue-Spotted Complex (includes A. laterale and A. unisexual that are A. laterale dependent)
b) Jefferson Complex (includes A. jeffersonianum and A. unisexual that are A. jeffersonianum dependent)

For the others, I have less experience with, but would suggest taking the same approach as above. In this manner, we can more accurately identify the "pure" species, and any researchers using the information can potentially make the assumption that there are unisexual species present as well. The only 100% identification we can make is the "pure" species in my opinion, as even a lab confirmed A. unisexual will confirm the presence of the associated "pure" species.

In any scenario, I am happy to help in any manner, and will modify my identifications going forward to align with whatever the outcome of this conversation is. I can also assist in combing back through identications to help align them to any new proposals should we get there.

Regards,

Chris

I like that idea. It seems practical enough. I guess there is the occasional area where Blue-spotted, Jefferson, or Unisex could occur, but I guess we should just leave that to genus. I mean, there are several good ideas here now, and I will be happy with any, my only hope is that it is as simple as possible, with still being scientifically usable data. I do think we need to keep the Blue-spotted taxon in case of an obvious male, but it may be just as simple to keep the “Unisex Complex” and we just have a clarification that it is for only true Unisex or this is for any and all Unisex types. I am happy though with how many ideas have came out of this discussion!

Got it. People can always use observations fields if they want to identify to a finer and more subjective level than iNat taxonomy allows.
How common are situations with all 3 where you can't tell if an individual is a "Tremblay's" or a "Silvery"?
If that's pretty rare then it sounds like it could work, the downside of this option is that I'm guessing it would prevent Ambystoma from being a "Complete Taxon". That may or may not be a dealbreaker, I don't understand the Complete Taxa stuff well enough.

@forth, if he does not respond, I might suggest Katy Greenwald at Eastern Michigan University as well.

Okay, so I think the main questions are:

1. Should Blue-spotted, Jefferson's, and Small-mouthed Salamander be species or complexes (without any lower taxa), and is the latter even allowed?

2. Should "hybrid unisexual complex" be a hybrid (as it is now), a complex (containing all 4/5 relevant species), or not exist at all?

For the first question, complexes would be more accurate given the practical ID situation but might not work with the way the taxonomy system is set up.

For the second question, I think it should be a complex because of all the ID confusion with having it as a hybrid, but I'm not sure if we can have complexes within a complex (in which case I'd say it shouldn't exist at all and should be merged with the genus).

I just would like to point out that in VA all Jefferson's are 100% Jefferson's and there is no chance of unisex cases (outside of a Tiger hybrid). I am less familiar with the Blue-spotted range extents, but I assume there are some places where there are no other species to hybridize with (maybe not). I may have miss understood something earlier, but I believe the species should stay on a species level, but probably the "Complex" taxon should be split.

We could retain both the pure species and a separate "species complex" for each. This would be more accurate than the single unisexual taxa dumping ground that we have now. In places without unisexuals, or when species is known (i.e., obvious males), this retains that level of precision.

Can we have geographic exclusion on taxa? As in, the species is unavailable around the Great Lakes where this is the issue, and everything defaults to the proposed complexes?
Here's why I ask: "Common" Gray Treefrog and Cope's Gray Treefrog are sympatric over a huge area, where they often can't be distinguished (unless they are calling). Currently iNat has species taxa for both as well as "Gray Treefrog Complex". In a place like Ontario, where only H. versicolor occurs, we can go to species reliably. Over much of the range of both species, most IDs should really be the complex. The result of having all taxa available in all places is that there are lots of "false" RG species-level records, as well as some records stuck in the complex in areas where only one species is present.

Some comments (and some added commentary from J. Bogart, where noted):

Current hybrid unisexual complex should be removed altogether. It is too broad, and not useful in its current state.
Setup Complex for Blue Spotted Salamander: Includes A. laterale and A. laterale-dependent. To be more precise, you could add the A. laterale-dependent species as A. 2-laterale jeffersonianum (J. Bogart: There are very few populations where two sperm donors are sympatric so it is pretty safe to identify the unisexuals as A. laterale 2-jeffersonianum or A. 2-laterale-jeffersonianum in allopatric areas of A. jeffersonianum or A. laterale.). @tysmith: We have areas where there are only Blue-Spotted (referring to your comment on areas with no unisexuals with jeffersonianum). I think that would still be covered under the complex correctly. I don't think there would necessarily be any harm in IDing in this manner.

Setup Complex for Jefferson's Salamander: Includes A. jeffersonianum and A. jeffersonianum-dependent. To be more precise, you could add the A. laterale-dependent species as A. laterale 2-jeffersonianum (J. Bogart: There are very few populations where two sperm donors are sympatric so it is pretty safe to identify the unisexuals as A. laterale 2-jeffersonianum or A. 2-laterale-jeffersonianum in allopatric areas of A. jeffersonianum or A. laterale.). @tysmith: Even where only pure Jefferson's exist, I think this would still be accurate.

For the remaining species of Ambystoma with unisexual presence (A. texanum, A. tigrinum and A. barbouri) the overlap is rare and mostly confined to small geographical locations, but it doesn't just exist on Peelee Island (J. Bogart: This becomes a problem in Ohio where you might get three sympatric species of sperm donors (A. jeffersonianum, A. texanum, A. tigrinum) in the same pond. Unisexual complex would probably be the only way to describe those unisexuals.). I would suggest in this scenario, we decide to do one of the following:

Make all of these species include a "complex", that would include their associated unisexuals similar to laterale and jeffersonianum and create an "A. unisexual-unidentified Complex". Anything that can't be related to a pure species, or one living sympatrically with multiple associated pure species could fall under this category. This could also include the instances of A. laterale and A. jeffersonianum living in close proximitely to each other as well).

Have no complexes for these species, but keep the "pure" species as identifiable. Create "A. unisexual-unidentified Complex" and ensure it instructs those using this as an identification that is includes only: A. laterale, A. jeffersonianum, A. texanum, A. tigrinum and/or A. barbouri that live sympatric with more than one of these species OR A. texanum, A. tigrinum and/or A. barbouri that are potentially unisexuals based on their geographic range.

I think looking at the above, the first option makes more sense. Further, it will also allow those who want to dig deeper, or have some constructive conversation, to look at the "A. unisexual-unidentified Complex" and know these are likely more complicated genome wise, and harder to identify, perhaps marking those areas as high interest areas for biologists/researchers etc. These identifications would probably be very common in very specific geographical ranges.
If the above changes can't be made (changing to complex for laterale/jeffersonianum and one of the above two options for the remainder) - I think the option of splitting these all up is the better option than leaving it as is. A. unisexual is not useful as it stands now, and certainly these aren't really hybrids in a typical sense. Right now, this is nothing more than a generalized dumping ground for identifications (worse yet, it doesn't help identify species that are endangered, such as those unisexuals that are dependent on jeffersonianum here in Ontario). Having a specific unisexual for each species would then be more accurate - but the problem is still there with that route: we now have two species, and two identifiers, with many areas that we can't properly determine which ID is the appropriate one.

For further context, I will make another post below this with the full response from J. Bogart. You can then see the quotes above in full context of what he was trying to say, in case I am inadvertently taking them out of context.

Sorry for the long one...

@upupa-epops is it possible to add user thomashossie to this discussion as well, for input? I believe he may have a slightly different take on this (and is a biologist, involved with some studies on Ambystoma from my understanding), which would be good to consider - but it will also allow him to comment on the discussion so far.

A. unisexual is not useful as it stands now, and certainly these aren't really hybrids in a typical sense. Right now, this is nothing more than a generalized dumping ground for identifications

@forth wouldn't this basically still be the case with both of your suggestions? It would be a bit less generalized, but still generalized and technically inaccurate?

The other currently possible option is what @pantherophis described:

the few observations that won't fall into one of those should be Ambystoma sp. - in those ambiguous cases, we can argue and speculate over photos all we want, but we really would need genetics to confirm whether it is unisexual or not, let alone which one. In super-rare cases where more is known, info can be added to notes or observation fields.

I don't know which is better. We seem to have a consensus on the other aspects though.

@upupa-epops The difference is, we have to choose between a unisexual ID, that spans all unisexual and possible unisexuals or the pure species. With the new arrangement, because there are other options, policing the unisexual "catch-all" is easier - and I imagine the description would further help people decide where to place an identification. It is there for a specific purpose, not a dumping ground for unknown identifications.

I just see a couple issues with the unisexual complex at the bottom of your taxonomy. The first is that it will likely cause confusion, with people identifying any unisexual as that complex when most should be identified as one of the other complexes. The other is that from what I can tell it's not accurate; as those various unisexuals aren't all of their closest relatives (although we can't get perfect accuracy with any method).

I don't see a good way to resolve them in with the current methods available though, so I think that's the best option we have at the moment. In the future if species group becomes an option, I'd suggest putting all of these taxa within a species group and losing that unisexual complex so those various unisexuals are just directly beneath species group level like A. barbouri and A. tigrinum would be.

personally, I don't think this is a great fit for the iNat taxonomy. Maybe tracking all of these breeds is better suited to observation fields maybe with an associated project?

You can see the 33 species of Ambystoma in our external taxonomic reference here. I don't think anyone is arguing that that those buckets perfectly capture the real world which is a gradient. But we don't want to entirely divorce the iNat taxonomy from the abstraction that is Linnean taxonomy for reasons ranging from making things easy to understand and use to transferable to other projects etc. I think the iNat amphib community would be ok with adding a complex or a hybrid or two to this genus, but I don't think we should consider multiple taxa made up of complex ratios of species as laid out above.

In the interest of resolving this flag - how about we present 3 options;
1) status quo leave Hybrid Ambystoma 'unisexual complex' as a oddly named hybrid
2) swap it into Ambystoma
3) alter the taxonomy with a single complex that strikes a compromise

if folks can agree in a month lets make the change to 2 or 3, but if not, I think we should stick with 1 and close this flag. Does that sound reasonable?

Anyway, I think we would be fine if we got rid of the "Complex A. Unisexual" complex in forth's system; all of the taxa in it are pretty rare and obscure from what I understand. If they're suspected, they could just be identified to genus level and an observation field could be used to track which one they're likely to be. That would just leave the 3 other easier to understand complexes.

Of your 3 options, I prefer option 3 as I described above: https://www.inaturalist.org/flags/79019#activity_comment_37e8b1e6-713a-4285-b9a4-988d5a0818ca
Ambiguous individuals or definitively unisexual individuals would be ID'd to complex level. Anything that looks like a Blue-spotted Salamander should be ID'd as a Blue-spotted (even though it could be Blue-spotted or Blue-spotted-dependent unisexual) and so on for the other species. There will still be regular disagreement about whether they should be ID'd to species or complex, but it's better than the current disagreement between species and "unisexual complex hybrid" which ends up pushing the observations back to genus level.
(I had suggested "Complex A. laterale" but this site gives a reason to call it "A. jeffersonium complex" instead, so I don't know)
I think this makes the most sense if unisexuals are to be considered a "biotype" of their host species (I don't think I understand what "biotype" means though).

I think that plan will make the available options even less accurate, and move iNaturalist records even further away from reality. Increasingly I find that people use iNaturalist as their #1 and often only reference on all things natural history, and treat it as authoritative even when it shouldn't be. Some people actually change the common names they use for things to match iNat, even when the name chosen here isn't the most widely used or most appropriate name. Simplicity in the name of accessibility isn't good if people come away with a less accurate understanding of reality, or less prepared to learn it.

So with that in mind, a great big Ambystoma complex will leave many with the impression that this genus is a big melting pot of hybrids, which it isn't.

The other problem with this is that it will leave a handful of people - basically the half dozen or so posting here - to "police" and "educate" the entire iNat universe on how to correctly identify that very-Jefferson-but-almost-certainly-unisexual salamander.

I think that A. laterale, A. jeffersonianum, and A. texanum should keep the specific complex structure proposed above, with "pure" and [whatever]-dependent unisexual as "subspecies". This will absorb the vast majority of unisexual observations. The others are rare enough that "Ambystoma sp." with observation fields will do. That's only three species with non-standard "complications", which isn't that complicated, really. Plus it's a lot more realistic.

I agree with pantherophis here. The changes suggested don't do anything really.

Take a slimmed down version of what I put earlier, maybe that will work as a compromise. This slots all the species in under the complex where necessary, if that even works in a hierarchy. The below is better than what we have now, I think, and won't involve creating all of those dependent unisexuals. Realistically, they would never have been used anyway, but were there to be accurate.

This way most ID's will fall under the complex, but the pure species still exist because males CAN be field/photo identified. Anything that is debatable can be ID'd at the Genus level.

Genus Ambystoma (as related to Unisexual Ambystoma)
----- species A. barbouri (Streamside Salamander)
----- species A. tigrinum (Tiger Salamander)
----- NEW - Complex A. laterale (Blue-spotted Salamander Complex)
---------- species A. laterale (Blue-spotted Salamander)
---------- (removed)
----- NEW - Complex A. jeffersonium (Jefferson Salamander Complex)
---------- species A. jeffersonianum (Jefferson Salamander)
---------- (removed)
----- NEW - Complex A. texanum (Small-mouthed Salamander Complex)
---------- species A. texanum (Small-mouthed Salamander)
---------- (removed)
----- (removed)
---------- (removed)
---------- (removed)
---------- (removed)
---------- (removed)
---------- (removed)
---------- (removed)
---------- (removed)

I like what @forth is presenting. I would leave "unisexual" out of it, and let those fall under the complex ID.

This way, the three species most frequently affected by this get a "complex", for the indeterminate individuals in areas where unisexuals are found, and the species level, for individuals that are known to be "pure" species, or that are from areas where unisexuals are not found. This works because we can sometimes confirm a "pure" animal visually, but it is almost impossible to truly confirm a unisexual without testing (giant A. laterale-type individuals may be the only exception). If a unisexual is somehow confirmed, that can easily be noted with observation fields on the complex level ID.

All other Ambystoma species are unaffected. The phylogenetic tree / Linnaean structure is thus mostly clean and simple, and there is no unisexual "dumping ground" taxon.

I would suggest that the three main "complexes" here should be "species" in the iNaturalist sense, i.e., the level suggested by AI, and Research Grade if confirmed through community ID. The true species-level IDs should be a "subspecies" in the iNaturalist sense: available for situations where the determination can be made, and generally only used by people who know what they're doing.

Having these complexes will also prevent a lot of arguing and downgrading of IDs. Currently well meaning people with an incomplete (or dated) understanding of the issue do this frequently. A few weeks ago someone made a case to me based on a (single) paper he just read on this - that was published in 1991. That kinda thing.

This was forth's original taxonomy, minus the extra unisexual complex containing the rarer and more complex unisexual taxa:

Genus Ambystoma (as related to Unisexual Ambystoma)
----- species A. barbouri (Streamside Salamander)
----- species A. tigrinum (Tiger Salamander)
----- Complex A. laterale (Blue-spotted Salamander Complex)
---------- species A. laterale (Blue-spotted Salamander)
---------- species A. (2/3/4)-laterale-jeffersonianum (Blue-spotted-dependent Unisexual Salamander)
----- Complex A. jeffersonium (Jefferson Salamander Complex)
---------- species A. jeffersonianum (Jefferson Salamander)
---------- species A. laterale-(1/2/3/4)-jeffersonianum (Jefferson-dependent Unisexual Salamander)
----- Complex A. texanum (Small-mouthed Salamander Complex)
---------- species A. texanum (Small-mouthed Salamander)
---------- species A. laterale-(1/2/3/4)-texanum (Small-mouthed-dependent Unisexual Salamander)

The numbers indicate the ploidy level and the slashes are a compromise representing that the taxon can be used for all the potential relevant ploidy levels for that kind of unisexual.

Maybe we can just have e.g. "A. jeffersonium-dependent" as the scientific names? Less accurate but easier to read and understand. Another option is how I wrote their names in this comment, which is how the Ontario government writes their scientific names, but also a bit inaccurate since it only has the most common possibilities: https://www.inaturalist.org/flags/79019#activity_comment_b0375b7c-2f6c-4587-a32f-12e4502e51a1

The unisexuals are not really hybrids, they are a species on their own right (possibly more than one). They break the rules of your typical taxonomy which is why this is a bit different than most discussions.

We are trying to make it as easy as possible for average users, but also for us who actively review the identifications, but an oversimplification isn't necessary what will make it easy.

@loarie I'm sorry you don't like the proposals. Unfortunately nature is messy, and iNaturalist needs to be more flexible here in order to be relevant. If there was ever a situation where the 250-year-old Linnaean structure is insufficient, this is it. Rather than shoehorning reality into an inadequate structure, why not create a structure that is both appropriate to reality and the practical needs of visual/field ID?

The pure species do not belong in a complex together, that is plainly wrong. They are separate entities, which can be reliably separated in the absence of unisexual lineages.

The unisexual lineages are not hybrids. They are independent entities, and are functionally species in ecological terms. They don't quite meet the official definition of species in the BSC because they don't exactly "produce like offspring", as their ploidy levels are flexible, and because they require spermatophores from different species to reproduce, even though they don't incorporate that genetic material.

This flag began because the current hybrid Ambystoma unisexual complex is inaccurate, incorrectly lumps all unisexual lineages into a single taxon, and becomes a dumping ground for any vaguely challenging IDs- and sometimes not challenging IDs - which results in the burying of potentially useful or significant observations of rare/endangered species.

Each of those three species can be reliably separated, but usually cannot be separated from its "respective" unisexual lineage. That doesn't mean a given species cannot be reliably separated from all unisexuals - just not from the one that is typically dependent upon it for spermatophores. Any phenotypical "Jefferson Salamander" may be a true A. jeffersonianum OR a "LJJ" (or LJJJ or LJJJJ) unisexual - but not an A. laterale or associated unisexual, nor an A. texanum or associated unisexual. This is why the three-complex split is the way to go.

OK, my personal opinion is that for amphibians its not worth the precedent and confusion incurred by adding these unisexual lineages to the taxonomy - e.g. "species A. laterale-(1/2/3/4)jeffersonianum (Jefferson-dependent Unisexual)"
so I'd be inclined to leave things the way they are (keep this single catch all Hybrid Ambystoma 'unisexual complex' taxon) or swap Hybrid Ambystoma 'unisexual complex' into genus and ID these taxa to genus Ambystoma and use Observation Fields to track them.

But this should be a community decision so I'm looping in some of the top Amphibian identifiers for more opinions.
@diogoprov, @martinmandak, @johngsalamander, @sullivanribbit, @amarzee, @alexanderr, @tom-kirschey-nabu, @lucareptile, @donchelu, @mario_yanez_munoz

The problem with a single complex is the problem we have right now - that anything inside the distributions of the unisexual lineages just gets drowned into that single melting pot, when it is possible to separate them 95%+ of the time. Personally I would rather not have any "complex" taxa rather than a single catch-all unisexual taxon.

In plants, sedges have every sub-generic "section" available for selection. In moths, there are multiple species-complex nodes for species that are hard to separate. Salamanders are vertebrates; "higher taxa" if you will. We're asking for three nodes for three species. Because of reality. We should not dumb down nature. It is what it is, and we are more than capable of recognizing it as such. If we aren't, what are we doing here?

Saying that, a pitch I made several zillion posts ago may be workable and even simpler, but may need to be rephrased. I had proposed making each of the three species in question:

-->"Ambystoma jeffersonianum or A. jeffersonianum-dependent unisexual"
-->"Ambystoma laterale or A. laterale-dependent unisexual"
-->"Ambystoma texanum or A. texanum-dependent unisexual"

This is the same as saying:

-->Complex Ambystoma jeffersonianum
-->Complex Ambystoma laterale
-->Complex Ambystoma texanum

...with no sub-taxa. Full species in an area with no unisexuals? No problem, it goes here. Indistinguishable adult in an area with unisexuals? No problem, it goes here. That is the simplest way.

Personally I favour having one sub-taxon for each of these, which would be the full species. No long taxa list of all the unisexual variants. That is slightly less simple, but not hard, and very manageable.

@upupa-epops The confusion you mention is coming from a conflation from a relatively recent theory about the origin of the unisexual lineages. All of the unisexuals may be monophyletic, but they are not a single entity. Most analyses estimate them to be hundreds of thousands, perhaps millions of years old. When they incorporate the "donor" spermatophore, the eggs undergo a process of meiotic reduction which cuts out the "donated" genetic material. Tetraploids and pentaploids happen (most are triploids) when this process doesn't occur "normally", which may be correlated with high incubation temperatures - loss of canopy cover from deforestation and climate change are implicated in increasing rates of these "abnomalities". On the other side of things, extreme meitoic reduction very, very rarely produces a diploid unisexual. The theory is that one of these diploids could incorporate a donor spermatophore from a different species and thus "found" a new lineage. This isn't an ongoing or dynamic process; this is what has been proposed as a mechanism for evolution and diversification in the unisexual lineages. That doesn't make them a single entity now; treating them as such would be like saying no species are real, and the genus is the lowest level of any taxon.

In practical, ecological terms: "pure" males have the ability to detect and avoid courtship with unisexual females to varying degrees. Males are better at avoiding unisexuals of the more "distant" unisexuals than the one that most resembles their own species - because each lineage is adapted to live with its "dependent" species. Also, in areas of overlap, "pure" species and their respective, "dependent" unisexuals are typically segregated in different breeding ponds or areas, in part to avoid larval competition for resources. Where this is not the case, anthropomorphic factors (habitat destruction/fragmentation) are typically responsible for "pooling" of species that would not be naturally "preferable". These are both part of the "ecological species concept" rationale for recognizing separate lineages as functional "species", as well as limiting factors that normally prevent the theorized "founding" of new lineages by diploid unisexuals.

I'm really not an expert in salamanders, but I'd also not use hybrid as a taxonomic rank

Option B is the best option in my mind for being able to identify these animals within existing taxonomic constraints.

Looking at the structure above, and how I laid it out, it may cause problems in identifications though.

The complex itself should be a species level ID, similar to how Hyla versicolor, Hyla chrysoscelis and Complex Hyla Versicolor work. We just need that option for each grouping.

It isn't perfect, but will help track and ID these much better than we currently do.

waiting for other amphibian identifiers to chime in here. My personal preference is not to further complicate the taxonomy and use observation fields for tracking these unisexual lineages

I like it. It is simple yet displays these three groups well enough to justify splitting them to me.

@wayne_fidler @kelton_ah @arborsphere @cramnaejvallieres @syrherp @twpierson
Tagging some of the top amphibian identifiers for around the Great Lakes area. What are your thoughts?

I was directed to this conversation after I mistakenly added the identification “Ambystoma 'unisexual complex'” to a “Jefferson” like salamander that had been identified to “Genus Ambystoma”. Without giving it enough thought it seemed inappropriate to leave the posting at the relatively vague generic level considering the size of that genus. Before going any further, I should also say I’m an amateur, general naturalist with a minimum grasp of the complicated genetics involved here. I also live in the same area as @upupa-epops and “only” have to deal with the jeffersonianum – laterale complication.
That said I like proposal B that allows for species specific identification where location rules out the possibility of complexes or where the photo is of such quality that the specimen can be identified to species by someone qualified to do so. At the same time, it allows for a level of identification finer than “genus” when the photo doesn’t allow separation of species and complex. As an amateur and, I suspect, for many Naturalists that aren’t up to speed on this issue, it would help educate them that this whole complex issue exists. Engaging the public is one of the prime purposes of the iNaturalist platform. This may create a bias to lump sightings into the “complex” category, but I suspect that is a safer situation given the complicated genetic involved and the fact this platform is based on photos. Moving a sighting to RG is already allowed by the option “Based on the evidence, can the Community Taxon still be confirmed or improved?” and so these could be caught in a search for anyone delving into the data.
That’s all I think I can add except to say I hope you can resolve this frustrating situation. Soon. And thank you all for your efforts on this.

my personal preference has always been for these to get ID'd to genus and for people to use observations fields to track complex issues relating to hybridization (ie swap Hybrid Ambystoma 'unisexual complex') into Genus Ambystoma) you can use observation fields to search for, map and do any other things related to filtering and organizing observations. But in my opinion these complex issues of hybridization are too difficult to try to shoehorn into the iNaturalist taxonomy which is hard to maintain.

if there's a strong need for some set of observations to be obscured then I guess we have to involve the tree and my preference would be to keep the modifications as simple as possible - e.g. single hybrid with obscuring status as we have here or single complex sitting above more than one species with obscuring status

Caleb is right - these are not hybrids, and they aren't "spontaneous events" in mixed species breeding ponds. (The various lineages almost always segregate into ponds with their obligate donor species.)

They aren't parthenogenic, either - they require a spermatophore from a "true species" male of their obligate donor species - although they don't incorporate the genetic material.

They technically aren't being called species because of the technical definitions of a species - they don't "produce like offspring" in the classical way, where the male is one of the true parents. They are basically gamete parasites. However, that is human semantics - we wrote the definitions that don't work here. In contemporary terms the various "taxa" are called "unisexual lineages", because they are "conservation units" with distinct populations and ecological significance.

In Ontario, the Jefferson-dependent unisexual lineage (with LJJ chromosomes, or in an antiquated approach "Silvery Salamander" or "Ambystoma platineum") has the same status (provincially and federally Endangered) as true, pure Jefferson Salamander (A. jeffersonianum). This is in part because they can't be separated readily in the field, but also because the Jeff-dependent unisexuals indicate the presence of pure A. jeffersonianum, by virtue of that reproductive obligation/parasitism. This is why I advocate for any system that either ties identifiable unisexuals to their obligate species, or one that allows different lineages to be identified separately. A Jefferson or a Jefferson-associated unisexual lineage (as well as the Smallmouth and Smallmouth-associated unisexuals) would be subject to legal protection in Ontario, including habitat protection (and different location-sensitivity defaults). An indeterminate, generic "unisexual" taxon or a default to just "Ambystoma" is not only less specific that we are actually able to be in many cases, it prevents rare lineages from being identified readily for conservation purposes. (Nevermind that is is taxonomically inaccurate.) The provincial agency responsible for tracking rare and endangered species primarily uses iNaturalist for reporting now.

@pantherophis I most definitely agree with your proposed system. @upupa-epops thanks for letting me know about this!

I'm supportive of https://docs.google.com/drawings/d/14Nzkv1lTVAYy5Vvp4gB3tbUSXuO9FVH6LHPMgZP9t7c/edit

@radbackedsalamander I'm curious what you mean by more restricted, for option B? I think it gives more flexibility since in your example you would be able to ID your salamanders as [laterale or laterale-dependent unisexual] as opposed to option A where your only choice is [unisexual, laterale, jeffersonianum, or texanum, etc.].

Regardless, going with option A with just the one complex as in your diagram we could still use observation fields to sort the visually identical [normal species and associated unisexual lineage] groups.

Alright, action items:
~ Change rank of this taxon to Species
~ Rename this taxon Ambystoma 'unisexual'
~ Create new taxon Complex Ambystoma 'unisexual' containing Unisexual Ambystoma and the other 5 species
~ Alert the people in charge of Canadian conservation statuses so they can make the complex auto-obscured in Ontario.

Does that sound good to everyone?

as for your second question I mean that the borders and ranges for the species dependent complexes are VERY fuzzy, we don't quite know where one begins and ends, and it's not as simple as appearance, many jefferson dependent unisexual look identical to laterale dependent unisexual

@radbackedsalamander ah yeah, in that case those would have to be identified to genus level (or subgenus now) using option B. At least here in Ontario in most cases (as far as I know) it's made easier by the fact that a specific pond or forest will only have Blue-spotted-type salamanders or Jefferson-type salamanders, not both.

In the iNaturalist taxonomy, complex is below subgenus so we would have to place the complex within a subgenus, and then the 5 species+unisexual have to be within the complex. That doesn't work if the species also have to be below their proper subgenera. @loarie
So either we get rid of the subgenera, or make a separate complex for each species that has an associated unisexual lineage (only 3 of the 5 species currently as far as I know) so that those complexes can be in their species' respective subgenera. (as I described above both systems are partially accurate genetically and ecologically from my understanding)

I created those subgenus following the taxonomy authority listed in the curator guide : Amphibiaweb. If I did a wrong thing I'm going to undo everything now. https://amphibiaweb.org/lists/Ambystomatidae.shtml

@loarie Can I make the change? In favor of: https://docs.google.com/drawings/d/14Nzkv1lTVAYy5Vvp4gB3tbUSXuO9FVH6LHPMgZP9t7c/edit

I would make Ambystoma 'unisexual complex' an invalid taxon and switch it over to Ambystom complex 'unisexual'.

👍

I also think an action should take place soon as this flag has been running for 5 years now @loarie

It already began but even sooner we will see a massive influx of observations from places like Vermont, Ontario, Maine, New York, etc.

Under that proposal presumably we would add an obscuring status to the complex parent of Blue-spotted Salamander which would obscure all Blue-spotted Salamanders.

To clarify, is it not possible to have the complex obscured and Blue-spotted open?

If obscuring statuses were added to Ambystoma for certain states, that would have the same impact on descending taxa than the complex approach.

Is it possible to auto-obscure based on observation fields? Is that what you mean here?

If you want me to add a new complex node that sits between Genus Ambystoma and the species here

Yes please, I think that would add significant clarity and utility for this group, even aside from concerns about obscuring.

To clarify, is it not possible to have the complex obscured and Blue-spotted open?

it is not, if a taxonomic ancestor is obscured all descendants are obscured

Is it possible to auto-obscure based on observation fields? Is that what you mean here?

thats not what I meant. I meant that adding your complex node won't improve the obscuring tradeoffs at all from the current situation without the complex node

Yes please, I think that would add significant clarity and utility for this group, even aside from concerns about obscuring.

I disagree. But I think you feel stronger than me so I'll make this change so we can finally close this flag

Ah I didn't realize that would force Blue-spotted to be obscured, that's unfortunate.

Thank you! I'll go work out the observation field.

At least this will be functional, if semantically wrong. The unisexuals aren't true hybrids, as has been said - but the structure of the "rank" levels here is good.

Obscuring Blue-spotted Salamanders isn't necessarily a bad thing - there's an argument to be made that all herp data should automatically be obscured, but that's a different topic.

I spend much of my time studying these animals, so I thought it might be helpful to provide some additional context and thoughts here.

In general, I prefer to treat all unisexuals in the genus Ambystoma as a single taxonomic unit, which is distinct from any of their contemporary host species (e.g., Blue-spotted and Jefferson salamanders are still distinct species from unisexuals). This is supported by genetic evidence from their mitochondrial DNA. The contemporary species with the most similar mitochondrial genome is the Streamside salamander. Molecular dating supports an early Pliocene origin for the unisexual linage (~5 million years ago), and shows that it is clearly disticnt from any contemporary species. The nuclear genome of unisexuals is ephemeral and changes over space and time depending on the host species that is currently available. Unisexuals vary in the number of chromosome complements (2-5 sets), and the combination of nuclear genomes they have (i.e., LLJ, LJJ). All unisexuals have at least one set of chromosomes from A. laterale, plus one or more from one of 4 other species. The amount and combination of nuclear DNA in the genome can change across generations of a single lineage thorough kleptogenesis, and genome replacement (i.e., an LJJ female can give birth to an LLJ offspring).

Finer classifications of unisexual in the genus Ambystoma might be useful for management (this is how COSEWIC and COSSARO in Canada operate), but this is more for pragmatic purposes given that confirming identity is hard/impossibly without genetics and some of the host species they live alongside (and are reproductively dependent on) are endangered. A given population of unisexuals at a specific site will be composed of a diverse assemblage of different unisexual forms that vary in ploidy and genotype (e.g., LJ, LLJ, LJJ, LLJJ, LLLJ etc.). Our contemporary understanding of the host ranges is also incomplete and may be outdated. Taken together, I don't see a good reason to apply finer classifications of unisexual Ambystoma. Personally I don't like the tendency of some users to identifying iNaturalist records as the host species when its possible, likely, or known that they are a unisexual. I'd rather have them left identified only to genus level. Virtually all unisexuals are female. Because males can be identified in the breeding season by their swollen cloaca, they can be reliably identified (as one of the host species) to species. It may be possible to identify a salamander as a unisexual in some cases if their phenotype falls well outside of the range of their host, but in many cases the ID is best left only to genus level without accompanying genetic evidence.

Here is some relevant material that might help inform this discussion:

Unisexual Salamanders in the Genus Ambystoma
https://bioone.org/journals/herpetologica/volume-75/issue-4/Herpetologica-D-19-00043.1/Unisexual-Salamanders-in-the-Genus-Ambystoma/10.1655/Herpetologica-D-19-00043.1.full
*This paper is a good overview of our contemporary understanding of these animals

A family study to examine clonal diversity in unisexual salamanders (genus Ambystoma)
https://cdnsciencepub.com/doi/full/10.1139/gen-2019-0034
*this papers outlines a recent development in understanding of the system (i.e., genome replacment)

Niche determinants in a salamander complex: Does hybridism or reproductive parasitism explain patterns of distribution?
https://esajournals.onlinelibrary.wiley.com/doi/full/10.1002/ecs2.3265

Time and time again: unisexual salamanders (genus Ambystoma) are the oldest unisexual vertebrates
https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3020632/

@freyathewolf haha yeah it's confusing. And yeah, so in my area we have Jefferson's and I know that there are unisexuals among them. Some individuals can be sexed as male based on having a swollen cloaca, and they can be identified as Jefferson's. Any other individuals that look like Jefferson's would be at the complex level unless you do a DNA test on them.

There's also a spot near me with a Blue-spotted Salamander population, and some individuals are the size of a Jefferson's Salamander (2-3x maximum Blue-spotted length), and I believe those can confidently be identified as unisexual.