Proposal (992) to South American Classification Committee

 

 

Treat Tyto furcata as a separate species from Barn Owl Tyto alba

 

 

Background:

 

Two recent proposals to NACC (2018-C-13 and 2022-B-6) have considered the taxonomic placement of various taxa within the Barn Owl (Tyto alba) complex. Comments on both NACC proposals and one submitted concurrently to SACC (#908) raised concerns about the lack of analysis of vocal differences among taxa.

 

Although pointed out in comments under the previous proposals, we highlight here a distinctive and prominent flight call associated with mate attraction that is uttered by New World Barn Owls and is absent in Old World Barn Owls. Based on this and concordant genetic data, we recommend adoption of New World Tyto furcata as a separate species from the Old World taxa. Work that might refine understanding of the Barn Owl complex both within the New World and separately in the Old World is discussed along with what is known about vocal and plumage differences. Genetic data presented in the previous proposals are included for the sake of completeness.

 

The cosmopolitan Barn Owl (Tyto alba) has a long and complex taxonomic history, with the American, African, southeast Asian, Australian, and many insular taxa being considered full species at various points. The current AOS taxonomy (AOU 1998) is largely based on Peters (1940) who lumped many previously recognized species under a cosmopolitan Tyto alba, with 34 then-recognized subspecies. When the AOU expanded coverage to include the West Indies and Middle America, T. glaucops (previously subsumed under T. alba by Peters 1940) was recognized as a separate species given its sympatry with T. a. pratincola (AOU 1983). More recently, some authors have opted to consider the American furcata clade and the southeast Asian + Australian javanica clade as two species separate from the alba clade of Europe and Africa (e.g., Gill et al. 2024). Additionally, three insular taxa from the Macaronesian islands are occasionally elevated to species level (Robb 2015), as are some insular taxa in the Indian Ocean and Indonesia. Many of these insular taxa are much darker than their mainland counterparts, including some with dark facial disks. These are all outside our area but highlight that species limits in the complex are highly dynamic, and that insular taxa especially are treated as full species by some authors.

 

For reference pertinent to this proposal, select taxa and subspecies groups (based on Clements 2023) along with their respective distributions are listed below:

 

• alba (Scopoli, 1769). Subspecies group (4 taxa) Europe, n. Africa, and Middle East east to Iran (hereafter alba ssp. group); the alba clade as a whole includes the alba ssp. group plus six other subspecies that occur on islands off Africa (5 taxa) and across sub-Saharan Africa (1 taxon, T. a. poensis), each regarded as a separate subspecies group by Clements (2023).

• javanica (Gmelin, 1788). Subspecies group (6 taxa) Pakistan east across s. Asia to Australia; also referred to as javanica clade.

• furcata (Temminck, 1827). In sensu stricto (s.s.) refers to T. a. furcata, a monotypic subspecies group, White-winged Barn Owl (Clements 2023), of Cuba, Isle of Pines, Cayman Islands, and Jamaica; elevated to species rank based on osteological differences by Suárez and Olson (2020); sometimes regarded as part of tuidara subspecies group. For this proposal, furcata clade or simply furcata refers to all 11 subspecies in the Americas, including tuidara group, currently classified under T. alba (sensu lato, s.l.) and proposed to be split as T. furcata.

• tuidara (J. E. Gray, 1827)*. Subspecies group (6 taxa) ranges from Canada to Tierra del Fuego. Type locality of tuidara is Brazil. [* see footnote on publication year]

• punctatissima (Gould & G. R. Gray, 1838). Galápagos.

• pratincola (Bonaparte, 1838). Mainland North America south to southern Mexico, recently to Hispaniola. Part of the tuidara subspecies group.

• glaucops (Kaup, 1852)*. Hispaniola. [* see footnote on publication year]

• insularis (Pelzeln, 1872). St. Vincent south to Grenada. With nigrescens grouped as Lesser Antilles Barn Owl (Clements 2023) or as a species (Suárez and Olson 2020); regarded as subspecies of T. glaucops by Bruce (1999) and Gill et al. (2024).

• nigrescens (Lawrence, 1878). Dominica. With insularis grouped as Lesser Antilles Barn Owl (Clements 2023) or as a subspecies under insularis (Suárez and Olson 2020); regarded as subspecies of T. glaucops by Bruce (1999) and Gill et al. (2024).

 

New information:

 

Vocalizations:

 

One of the primary issues raised by committee members in previous proposals is the lack of analysis of vocalizations in the Barn Owl complex. Although no formal analysis is yet published, we think that the qualitative analysis provided here is sufficient to elevate the furcata clade to species rank. Across the genus Tyto and within the Barn Owl complex there are a wide array of both vocal and mechanical sounds. Here we focus on the context of vocalizations associated with breeding, which is also the time when these owls are most vocal. Two specific types of vocalizations are defined below: Screech and kleak-kleak.

 

  Screech: Categorized as either courtship or perennial (Robb 2015). Recordings below are from https://soundapproach.co.uk/species/common-barn-owl/

 

(1) Courtship screech: Used by males of alba clade (here specifically alba ssp. group), typically given when perched but also in flight. Courtship screech in addition to perched context is also longer and with shorter gaps between calls compared to the perennial screech. Existence and context of this courtship screech is unknown in the furcata clade (G. Vyn fide Robb 2015). Notably, none of us has ever experienced a bird of the furcata clade screech from a perch. This needs further investigation.

 


Spectrogram of courtship screech by T. a. alba (Robb 2015)

 

(2) Perennial screech: Used by both sexes, uttered in flight and less often from perch in alba clade but perhaps never (or rarely?) given from perch in furcata clade. Further investigation between the perennial screech of the alba clade and the flight calls of the furcata clade is needed, especially in the context of whether the call is uttered when flying or perched.

 

• kleak-kleak (Vyn 2006): Given in flight by furcata clade, perhaps most often used by unpaired males (Gerrit Vyn pers comm fide M. Robb) or males in vicinity of nest (Marti et al. 2020); presumed to have an important role in mate attraction. Absent in both alba and javanica clades. Sometimes categorized under terms like cackles, chirrups, or twitters.

 

Screenshot 2024-01-03 at 3.26.58 PM.png

 

 

 

 

 

 

 

 

Spectrogram of kleak-kleak by T. a. pratincola (Vyn 2006)

The screech (or scream in Marti et al. 2020) is the best-known vocalization. The kleak-kleak call was described under “chirrups and twitters” in Marti et al. (2020). We note that much published information on vocalizations draws on Old World studies. Thus, it is important to heed the warning in Marti et al. (2020):

 

“Other than anecdotal notes, only unpublished information is available on vocalizations by the North American race (E. McLean and B. Colvin pers. comm.). Some of the calls described […] have not been positively documented for the North American race.”

 

Indeed, much of the behavioral context and sounds ascribed to Barn Owls in the Americas is adopted from Old World literature. Our summary here is guided in large part by “The Sound Approach” (Robb 2015), with especially helpful material published by that author on Barn Owls of the alba ssp. group here. One of us (O.J.) perused the sonograms of all available Old World recordings on Xeno-canto (1,080 alba clade and 62 javanica clade), plus a large selection in the Macaulay Library. We found no examples of kleak-kleak in either alba or javanica clades.

 

From listening to recordings of many Tyto species, including glaucops and various Masked/Grass owls it is clear that the loud screech call is fairly conserved across the genus. There is some variation in length of the call among species, and some have a whistled quality, but there is also much intra-taxon variation in call length, perhaps related to whether these are courtship or territorial, perennial screeches.

 

Typical screech calls of the three clades are given below. For javanica and alba clades, the screech tends to fade out and fall in pitch at the end of the calls, unlike furcata clade which ends more abruptly and rises slightly at the end:

 

alba: https://macaulaylibrary.org/asset/301733691

javanica: https://macaulaylibrary.org/asset/117266311 and https://macaulaylibrary.org/asset/271631421

furcata: https://macaulaylibrary.org/asset/50147

 

European birds (alba) do tend to give longer screech calls than furcata, while javanica are generally shorter but with a subtly different quality than furcata. However, alba and javanica commonly have a harsh whistled quality to the notes:

 

alba: https://macaulaylibrary.org/asset/235237551 and https://macaulaylibrary.org/asset/367445881

javanica: https://macaulaylibrary.org/asset/273379781

 

Here is an exceptionally long screech call from furcata: https://macaulaylibrary.org/asset/425012341

 

Non-screech calls, when present, seem quite different among species. The Australian Masked-Owl (T. novaehollandiae) utters a call called a cackle that is said to be given in courtship display flights by males circling over breeding territory (Higgins 1999, page 919). An example of that cackle call is here (https://macaulaylibrary.org/asset/539506871) and seems analogous to the kleak-kleak call of furcata. Likewise, analogous vocalizations exist in the two grass owls, T. capensis and T. longimembris (Robb 2015).

 

The kleak-kleak call of furcata is present across its range, with recordings from California, Florida, and Brazil. Here are a few examples:

 

https://macaulaylibrary.org/asset/172455681

https://macaulaylibrary.org/asset/245778421

https://macaulaylibrary.org/asset/554918181

 

Critically, this “kleak” call appears to be entirely absent from both alba and javanica according to The Sound Approach and our own perusal of recordings. Robb (2015), quoting Gerrit Vyn (pers. comm.), says that “unpaired males use this call most often…so it must have an important role in mate attraction.” Marti et al. (2020) also report that males give the “kleak” call in the vicinity of the nest, soon after leaving the daytime roost, and when approaching with food deliveries. Given that analogous calls exist in T. novaehollandiae and other Tyto, we suspect it has been lost in alba and javanica. Regardless, in our view this is a diagnostic vocal difference between the clades.

 

In our personal experience, this “kleak” call is nearly always given in flight. For example, JLD recently witnessed (summer 2023) one bird giving the kleak-kleak call in fluttery flight almost nonstop for a few minutes as it circled a lit up area near a known nest. The only mention that we can find regarding the “kleak” call for alba is Bunn et al. (1982), who say that it is reportedly uncommon in Britain. This contradicts Robb (2015) who has extensive experience with the alba ssp. group in Portugal and elsewhere. Despite fairly exhaustive searches of databases online we are unable to find any recordings of this vocalization from anywhere in the Old World. This reference of the kleak call in Britain appears anecdotal and could refer to another call that Bunn (1977) called the kit-kit call.

 

We feel it worth mentioning that no North American Field Guide or popular book on owls, including König et al. (1999) and Weidensaul (2015), mention the kleak-kleak call or its context in display. How did the birding community miss this characteristic sound of New World birds? The one source that does have it is Marti (1992), but none of us picked this up.

 

Genetics:

 

A paper by Uva et al. (2018) analyzed two nuclear and five mitochondrial loci to estimate a phylogeny of Tytonidae. This paper was included in the 2018-C-13 NACC proposal, and the proposal included a haplotype map that was based on a single mitochondrial gene but did not include the phylogeny that was based on a greater set of genes. That proposal did include the phylogeny from earlier work by Alibadian et al. (2016) that was based on slightly fewer genes and many fewer taxa. Although comments from many committee members considered the genetic evidence inconclusive on its own, we include it in the current proposal for the sake of completeness. Relevant figures from Alibadian et al. (2016) and Uva et al. (2018) are reproduced below.

 

 

Phylogeny from Alibadian et al. (2016):

 


 

Sampling map, haplotype network, and phylogeny from Uva et al. (2018):

 

 

Picture 2

 

 

A map of the world with different colored dots

Description automatically generated

Picture 3

 

Based on the genetic data, the current circumscription of Tyto alba comprises three major clades: alba, furcata, and javanica, with the former two being sisters. Uva et al. (2018) advocated elevating both furcata and javanica to species rank. Whether the alba and javanica clades should be treated as species is outside our purview and perhaps should await potential future contact (see Additional Considerations, below).

 

A few issues arise. First, Tyto glaucops is embedded within the furcata clade, being sister to punctatissima of the Galápagos, the two in turn being sister to the rest of the furcata clade. However, Uva et al. (2018) note that “given the poor node support (0.77 PP/66 BS) putative genetic distinctiveness of Caribbean and Pacific populations needs further confirmation” and we agree with that assessment. Regardless, the species status of punctatissima should be left to SACC.

 

Another issue is that two dark taxa of the Lesser Antilles, nigrescens and insularis, were not sampled by Uva et al. (2018); these taxa have been considered as subspecies of Tyto glaucops or as their own polytypic species (Suárez and Olson 2020). Given the lack of genetic and vocal information on these taxa, we think it best to leave them as subspecies of alba (or furcata if this proposal is adopted) for now, pending further study. Also see Additional Considerations, below, regarding anecdotal information on Barn Owl calls heard on Grenada where insularis occurs.

 

A recent paper on Barn Owls of the West Indies by Suárez and Olson (2020) was the basis for NACC proposal 2022-B-6, which did not pass but focused on the species status of glaucops, nigrescens, and insularis plus some extinct forms. Suárez and Olson (2020) analyzed osteological data from extinct and extant Caribbean Tyto. They elevated the taxon T. a. furcata Temminck 1827 of Cuba, the Cayman Islands, and Jamaica to species rank, leaving tuidara J. E. Gray 1827 as the name for the American mainland species. However, their osteological comparisons were to alba of Europe rather than to pratincola of the United States, so the question of species rank for furcata s.s. is unresolved. With regard to the priority of furcata for American Barn Owls over tuidara if split from alba, see footnote establishing that furcata has priority. Also note that furcata s.s. is considered a separate subspecies group by Clements (2023) based on the paler white plumage, especially of the wings. If, in the future, furcata s.s. is elevated to species rank, then the name for the remaining American barn owls would be tuidara Gray 1827. We note that Uva et al. (2018) sampled one individual that they labeled as furcata s.s. (sample number IPMB 20859), but no list of detailed sample localities is given in the paper or supplementary data and there is no dot from Cuba, Jamaica, or the Cayman Islands (the distribution of furcata) on their sampling map; moreover, we do not recognize the museum acronym and were unable to find a relevant record on VertNet or GBIF. Thus, it is unclear to us if true furcata was sampled by Uva et al. (2018). Although it would be the nominate taxon of the American clade, we think it extremely unlikely that it would be more closely related to Old World taxa than to mainland North American taxa, so it should not affect the separation of furcata clade from alba + javanica clades. It may have implications for the taxonomy of other Caribbean Tyto, however, if those are elevated to species rank in the future.

 

Plumage coloration:

 

Romano et al. (2019, Figure 2 reproduced here) showed that plumage coloration appears closely tied to rainfall and temperature. As can be seen in their maps, overall plumage coloration and spot size are not drastically different between the three clades (furcata, alba, and javanica). Nevertheless, the plumage and size of several taxa within the Americas do appear quite distinctive, e.g. punctatissima of the Galápagos, bargei of Curaçao, and insularis/nigrescens of the Lesser Antilles. Indeed, Ridgway (1914) separated these taxa and glaucops from furcata s.s. and the remaining mainland American Barn Owl taxa based on non-overlapping size, among other characters. Although not part of this proposal, we would not be surprised if more detailed studies suggest splitting more of these insular New World taxa. Interestingly, Ridgway (1914) noted that bargei is similar to nominate alba of Europe in coloration but is much smaller. We note that Uva et al. (2018) sampled bargei and found it nested within the furcata clade.

 

Picture 4

Additional considerations:

 

We currently consider Tyto glaucops unambiguously a separate species from T. alba s.l. based on sympatry on Hispaniola. Earlier authors, however, considered glaucops conspecific with alba s.l. (e.g., Hartert 1929, Peters 1940). On Hispaniola, T. alba either colonized sometime after 1930 or was overlooked before that (Keith et al. 2003). The source population is thought likely to have been pratincola from the mainland or Bahamas (Marti et al. 2020). Species limits considered by earlier authors were based on the same issues that we are dealing with currently, namely plumage and vocalization differences (but now also genetic) among allopatric insular populations. However, once colonization by alba s.l. occurred, it became clear that glaucops and alba s.l. were distinct species, a treatment followed ever since.

 

We listened to available recordings of glaucops (of which there are few, see examples below) and were not struck by major differences from furcata, which raises additional questions. If furcata and glaucops are sympatric, how are these being maintained as separate species despite the lack of described vocal differences? The screech call of glaucops seems a bit longer and more descending compared to furcata, which is interesting. If that is the case, then there are some minor vocal differences in the screech. A similar kleak call to the furcata clade is uttered by glaucops and could be taken as further evidence that this is a major character separating all New World barn owls (broadly speaking) from Old World barn owls. This, then, would be more evidence for splitting furcata. On the other hand, if plumage differences are keeping glaucops and furcata separate, how does that fit into our understanding of species limits in the genus given that plumage seems to covary with all sorts of things not related to species limits? Maybe the fast evolution of plumage in the genus allows for occasional evolution of drastically different-looking species?

 

Here is an example of the cackling kleak call by glaucops:

https://macaulaylibrary.org/asset/175146681

 

And recordings of its screech:

https://macaulaylibrary.org/asset/163149861

https://macaulaylibrary.org/asset/180725

 

We note in passing that Alvaro Jaramillo suggested that Barn Owls Grenada (insularis) gave vocalizations “much more like Ashy-faced Owl than Barn Owl” (Norton et al. 2005, page 512). Jaramillo’s analysis is repeated by Wiley (2021, page 209), who himself reviews the taxonomic history of Barn Owls in the eastern Greater Antilles through the Lesser Antilles. Note that east and south of glaucops on Hispaniola (and formerly Puerto Rico; Suárez and Olson 2020), Barn Owls occur on Dominica (nigrescens) and then on St. Vincent, some islands in the Grenadines, and south to Grenada (insularis), with no confirmed records for intervening Martinique and St. Lucia (Wiley 2021). To our ears, the calls of T. glaucops do not sound that different from the furcata clade so opining about calls of insularis on Grenada might be difficult without careful analysis. Some recordings of insularis sound similar to vocalizations of mainland furcata (https://macaulaylibrary.org/asset/541151851) but others do sound quite different and rather like some recordings of alba s.s. (https://macaulaylibrary.org/asset/317964701).

 

Given that the node separating glaucops/punctatissima from furcata is 1.75 Ma (Uva et al. 2018), it seems to us a reasonable yardstick extension to consider the much older splits of alba and javanica as different species from furcata (javanica vs. alba/furcata is 6.25 Ma, alba vs. furcata is 4.35 Ma). The alternative here is that glaucops/punctatissima are a recent offshoot from furcata that (unambiguously in glaucops) evolved reproductive isolation, while furcata and alba have not. We think that this is unlikely given that glaucops seems to have evolved reproductive isolation despite limited or no differences in vocalizations, while the limited vocal data we have indicate very distinct vocalizations between furcata and alba + javanica (primarily the lack of a “kleak” call in the latter as well as existence of courtship screech in at least alba ssp. group in alba clade). We also note that the node uniting glaucops and punctatissima has lower support (0.77 posterior probability/66% bootstrap) than most other nodes in that part of the tree, so the furcata clade may not be paraphyletic with broader genomic sampling. The node separating glaucops and punctatissima is 0.44 Ma. Uva et al. (2018) did not provide confidence intervals on these node date estimates.

 

There is also limited evidence that at least furcata and javanica are reproductively isolated. Populations from each of those clades were introduced onto Lord Howe Island to control rats: T. a. delicatula from the Australian mainland in 1923, and T. a. pratincola from the San Diego Zoo in 1927 (Hindwood 1940). Birds from these two taxa were not known to interbreed, and this was taken as evidence that the two should not be considered the same species (Bruce 1999). The only Barn Owl specimens collected from Lord Howe are of the Australian population, and no Barn Owls are known to have persisted past the mid 1980s (McAllan et al. 2004). It is presumed the American birds died out soon after introduction. This contact between the javanica and furcata clades could suggest that assortative mating was taking place, but the period of sympatry was brief compared to the longer period of sympatry between pratincola and T. glaucops on Hispaniola. We do note that javanica is the more distant clade in the phylogeny and does not provide direct evidence of species rank for the alba clade versus the furcata clade. However, it does suggest that multiple species exist within the cosmopolitan Barn Owl.

Finally, the International Ornithologists’ Union Working Group on Avian Checklists (WGAC) has recently split Barn Owl into three species, elevating the javanica and alba clades in addition to furcata. Although recognizing two Old World species is outside our purview, support for this is based on  morphological differences (Dick Schodde fide T. Chesser) and genetic evidence showing that the javanica and alba clades are not sisters (Uva at al. 2018). It is important to note, however, that Barn Owls have expanded east across much of Iran starting in the 1990s (Osaei et al. 2007). Prior to this, when the species was rare in Iran, specimens were ascribed to T. a. erlangeri of the alba clade (Vaurie 1965). The easternmost record in Iran (subspecies unknown) is at Bam, Kerman Province (Osaei et al. 2007), which is 900 kilometers (560 miles) west of the western limit of T. a. stertens of the javanica clade on the Indus Plain, eastern Pakistan. Thus, future contact between javanica and alba is possible, and further research would help to elucidate whether reproductive isolating mechanisms such as vocalizations exist to maintain species-level differences. Nevertheless, we think it is worth separately considering elevating javanica to species rank to align with this global checklist.

 

Recommendation:

 

We recommend splitting Tyto alba (Scopoli, 1769) into at least two species to recognize the vocal and genetic distinctiveness of New World taxa as American Barn Owl, Tyto furcata (Temminck, 1827). If the javanica and alba clades are considered conspecific, then Common Barn Owl is typically used for the Old World taxa.

 

English names: American Barn Owl is in wide usage by authorities that split furcata from alba, and we recommend that it be adopted. American Barn Owl was used by Ridgway (1914) for pratincola. Because of the possibility of paraphyly with glaucops and various other taxa embedded within javanica, we think that “Barn Owl” should not be hyphenated unless there is interest in renaming glaucops to “Ashy-faced Barn-Owl”. “American” in this context refers to the two continents on which this species occurs.

 

If the javanica and alba clades are retained as conspecific, then Common Barn Owl is typically used for the Old World taxa. However, the IOC (Gill et al. 2024) recognizes javanica and alba as Eastern Barn Owl and Western Barn Owl, respectively. Clements (2023) uses Eastern Barn Owl, Eurasian Barn Owl, and American Barn Owl for the subspecies groups. These English names are not ideal and potentially misleading (e.g. “Eurasian” occurs in Africa, and “Eastern” and “Western” could be confused with eastern and western North America). Therefore, consideration or solicitation of alternative names for the Old World taxa is merited.

 

Acknowledgments and Footnotes:

 

David Donsker helped research publication dates for relevant taxa. Alan Peterson’s Zoonomen.net website provided notes and insights on the publication dates of original descriptions.

 

* tuidara (J. E. Gray, 1827): This name was published at earliest 1 December 1827. Gill et al. (2024), among others, use 1828, whereas Bruce (1999) and Peters (1940), for example, use 1829. The name Tuidara Owl of John Edward Gray appeared in part 14 of Griffith’s Animal Kingdom, and this part was published 1 December 1827 (see table here). Temminck’s “Strix furcata” was published 30 June 1827 in livraison 73, plate 432 of the “Nouveau recueil de planches coloriées” (see table here and would therefore have priority regardless of the confusing dates ascribed to tuidara. The date of 1827 was used for tuidara by Suárez and Olson (2020), presumably based Cowan (1969). We use it here for the same reason.

 

* glaucops (Kaup, 1852): We found conflicting dates for this publication. The fourth edition of Howard and Moore checklist (Dickinson and Remsen 2013), Bruce (1999), and AOU (1998) use 1853. Gill et al. (2024), Peters (1940), and older publications use 1852. Note that Murray Bruce later agreed 1852 is the correct date (see notes here).

 

Literature cited:

 

Alibadian M., N. Alaei-Kakhki, O. Mirshamsi, V. Nijman, and A. Roulin. (2016). Phylogeny, biogeography, and diversification of barn owls (Aves: Strigiformes). Biological Journal of the Linnean Society 119, 904–918.

American Ornithologists' Union. (1983). Check-list of North American birds. 6th ed. Am. Ornithol. Union, Washington, D.C.

American Ornithologists' Union. (1998). Check-list of North American birds. 7th edition. Washington, D.C.: American Ornithologists' Union.

Bruce, M.D. (1999). Family Tytonidae (Barn-owls. In Handbook of the Birds of the World, Vol. 5), pp. 34–75 (J. del Hoyo, A. Elliott, and J. Sargatal, Editors). Lynx Editions, Barcelona.

Bunn, D. S. (1977). [Letters] The voice of the Barn Owl. British Birds 70(4):171.

Clements, J. F., P. C. Rasmussen, T. S. Schulenberg, M. J. Iliff, T. A. Fredericks, J. A. Gerbracht, D. Lepage, A. Spencer, S. M. Billerman, B. L. Sullivan, and C. L. Wood. (2023). The eBird/Clements checklist of Birds of the World: v2023.

Cowan, C.F. (1969). Notes on Griffith's Animal Kingdom of Cuvier (1824-1835). Journal of the Society for the Bibliography of Natural History 5:137–140.

Gill, F., D. Donsker, and P. Rasmussen (Eds). (2024). IOC World Bird List (v 14.1). doi 10.14344/IOC.ML.14.1

Hartert, E. (1929). On various forms of the genus Tyto. Novitates Zoologicae 35:93–104.

Higgins, P., Ed. (1999). Handbook of Australian, New Zealand and Antarctic Birds. Volume 4. Parrots to Dollarbird. Oxford University Press, Melbourne, Australia.

Hindwood, K. A. The birds of Lord Howe Island. Emu 40:1–86.

Keith, A. R., J. W. Wiley, S. C. Latta and J. A. Ottenwalder. (2003). The Birds of Hispaniola: Haiti and the Dominican Republic. British Ornithologists’ Union Check-list No. 21. The Natural History Museum, Tring.

König, C., F. Weick, and J.-H. Becking (1999). Owls. A Guide to the Owls of the World. Pica Press, Robertsbridge, UK.

Marti, C. D. (1992). Barn Owl. In The Birds of North America, No. 1 (A. Poole, P. Stettenheim, and F. Gill, Eds.). The Academy of Natural Sciences, Philadelphia; The American Ornithologists' Union, Washington D.C. https://doi.org/10.2173/tbna.1.p

Marti, C. D., A. F. Poole, L. R. Bevier, M.D. Bruce, D. A. Christie, G. M. Kirwan, and J. S. Marks (2020). Barn Owl (Tyto alba), version 1.0. In Birds of the World (S. M. Billerman, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.brnowl.01

McAllan, I. A. W., B. R. Curtis, I. Hutton, and R. M. Cooper. (2004). The birds of the Lord Howe Island group: a review of records. Australian Field Ornithology 21 (Supplement) 1–82.

Norton, R. L., A. White, and A. Dobson. (2005). The Spring Migration: West Indies & Bermuda Region. North American Birds 59(3):511–513.

Osaei, A., A. Khaleghizadeh, and M. E. Sehhatisabet. (2007). Range extension of the Barn Owl Tyto alba in Iran. Podoces 2(2):106–112.

Peters, J. L. (1940). Check-list of the birds of the world. Volume IV. Harvard University Press, Cambridge, Mass.

Ridgway, R. (1914). Birds of North and Middle America. Bulletin of the United States National Museum 50, part VI.

Robb, M. & The Sound Approach. (2015). Undiscovered owls, A Sound Approach guide. The Sound Approach. (web version available here: https://soundapproach.co.uk/undiscovered-owls-web-book/)

Romano, A., R. Séchaud, A. H. Hirzel, and A. Roulin. (2019). Climatedriven convergent evolution of plumage colour in a cosmopolitan bird. Global Ecology and Biogeography 28:496–507.

Suárez, W., and S. L. Olson. (2020). Systematics and distribution of the living and fossil small barn owls of the West Indies (Aves: Strigiformes). Zootaxa 4830:544–564. https://doi.org/10.11646/zootaxa.4830.3.4

Uva V., M. Päckert, A. Cibois, L. Fumagalli, and A. Roulin. (2018). Comprehensive molecular phylogeny of barn owls and relatives (Family: Tytonidae), and their six major Pleistocene radiations. Molecular Phylogenetics and Evolution 125 (2018):127–137. https://doi.org/10.1016/j.ympev.2018.03.013

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Louis Bevier, Carla Cicero, Jon L. Dunn, Rosa Alicia Jiménez, and Oscar Johnson

February 2024

 

 

 

Comments from Jaramillo: “YES to this proposal. I have been listening to and recording a few Barn Owls over my house recently, and had the same general thought. How could we have missed that the clicking call is the important vocalization in reproductively related display, not the screech. I have been listening to a pair over my house that sometimes displays overhead and actually seem to have an aerial "dance" of sorts where they cartwheel over each other, although it is hard to see as it is pretty dark even with the town lights. But the birds are clicking at each other, and sometimes one of them screeches, but the clicking call is the key.

 

https://nam04.safelinks.protection.outlook.com/?url=https%3A%2F%2Febird.org%2Fchecklist%2FS154888905&data=05%7C02%7Cnajames%40lsu.edu%7Cbd347e4679644ac5c0b408dc230e4a90%7C2d4dad3f50ae47d983a09ae2b1f466f8%7C0%7C0%7C638423793896727718%7CUnknown%7CTWFpbGZsb3d8eyJWIjoiMC4wLjAwMDAiLCJQIjoiV2luMzIiLCJBTiI6Ik1haWwiLCJXVCI6Mn0%3D%7C40000%7C%7C%7C&sdata=7fBhJfV%2FZiFGusTPiRu%2Bga3tatlUBqKCRiqb1gQ5mJI%3D&reserved=0

 

“Regarding Ashy-faced, Lesser Antillean and Galapagos birds. The possible genetic relationship between Galapagos and Antillean birds is not surprising to me. This pattern happens over and over again in multiple taxa. There is a Galapagos-Caribbean connection that has not been studied. Galapagos mockers are sister to Bahama Mockingbird, Galapagos finches are sister to the mainly Caribbean closed nest tanager radiation (the "quits"), even the flamingo is a connection. I worked with a Cuban spider expert who mentioned that Cuban spiders had some relatives in Galapagos. The fact that this has not been looked at more closely is interesting to me. I assume that there are some islands that are gone that connected the two before the Panama land bridge closed. In any case, I bet that in the future there is a confirmation of an Antillean/Galapagos Barn Owl clade.”

 

Comments from Robbins: “YES. What an in-depth proposal! Kudos to Bevier et al. for the extensive perspective on this.  Their proposal along with Alvaro's comments about the clicking call seem to underscore the importance of that vocalization: present in New World and apparently absent in Old World taxa. That coupled with the genetic data support recognizing New World furcata as a separate species from Old World alba. I vote yes in support of this split.”

 

Comment from Andrew Spencer: “Pieplow (2017) does mention the voc (which he calls "chitter") and does say it can be given by pairs in courtship.”

 

Comments from Niels Krabbe (voting for Elisa Bonaccorso): “YES. Back in the late 1960s I banded a brood of three Barn Owl chicks and had the luck to have all three recovered. One had flown 10 km SW, another 150 km N, and the third over 100 km E. Even though adult Barn Owls can be among the most resident birds and spend their entire life inside a single barn, young birds have a great capacity for dispersal, so I was fine with its worldwide distribution. As much as I hate to see another cosmopolitan go (there seems to be only Peregrine left), I see from the extensive, primarily mitochondrial comparisons by Uva et al. (2018) that the division of the Common Barn Owl group into three major clades is well-supported. The absence of the "kleak-kleak" sound in both alba and javanica clades and its presumed importance in mate attraction, as mentioned above, indeed supports species rank for the furcata clade. I was a bit surprised to find that several Xeno-Canto recordings of screeches by furcata were of perched birds, but far the most were in flight. Personally, I have only heard the short perennial screech, and only from flying birds. I was flabbergasted, when I first experienced a bird flying back and forth while giving the “kleak-kleak” call, which I had never heard in Europe.

“The proposal is elaborate and thorough, and I can find nothing to add. As stated, it is puzzling, that two Tyto species with quite similar vocalizations are both widespread on Hispaniola without interbreeding, and, also as stated, it would be interesting to know if the dark forms nigrescens and insularis (Lesser Antilles), belong with glaucops or furcata, but this is all outside the scope of SACC. As also discussed above, the mitochondrial similarity between glaucops and the Galapagos punctatissima has low support and needs to be substantiated by in-depth studies.

 

“I vote YES for ranking American Barn Owl as a species Tyto furcata.”

 

Comments from Areta: “YES. I was already convinced in https://www.museum.lsu.edu/~Remsen/SACCprop908.htm that there was overwhelming evidence for this split, and some unanswered questions on other Tyto. It is not that the "kleak-kleak" went unnoticed to Neotropical ornithologists, but it is the lack of it in the Old World that we (or I at least) were not aware of. Nice proposal, very thorough and clear.”

 

Comments from Stiles: “YES. Genetics and morphology found previously to support this split, but a thorough study of all available recordings now clearly demonstrates an important difference in vocalizations that tips the burden of proof onto those who favor maintaining the single-species status quo.”