Proposal (1044) to South American Classification Committee

 

 

Treat Formicarius destructus as a separate species from F. nigricapillus

 

 

Effect on SACC list

If this proposal passes, it will result in the elevation of the taxon destructus to full species, resulting in a monotypic F. nigricapillus and a monotypic F. destructus.

 

Background information

The current SACC note reads”

 

"1c. Areta & Benítez Saldívar (2025) provided vocal evidence for treating South American destructus as separate species.  SACC proposal needed."

 

Areta & Benítez Saldívar (2025) summarized the situation as follows:

 

The Black-headed Antthrush (Formicarius nigricapillus) includes two allopatric subspecies: nominotypic nigricapillus in Costa Rica and Panama, and destructus in Colombia and Ecuador (Ridgway 1893; Hartert 1898; Wetmore 1972; Krabbe and Schulenberg 2003, 2020). Although the taxon nigricapillus was originally described as a full species by Ridgway (1893) and treated as such by some authors (Chapman 1917, Cory & Hellmayr 1924), it was often considered as a subspecies of the Black-faced Antthrush (F. analis) (Hartert 1902, Ridgway 1911). Conversely, the taxon destructus was originally described as a subspecies of F. analis by Hartert (1898) and was subsequently either considered as such (Hartert 1902), rarely afforded species-level status (Salvadori and Festa 1899; Howell and Dyer 2022), or most often considered as a subspecies of F. nigricapillus (Chapman 1917, Cory & Hellmayr 1924, Wetmore 1972, Krabbe & Schulenberg 2003).

 

New information

In Areta & Benítez Saldívar (2025), we assessed species limits in F. nigricapillus using vocal, plumage, and morphometric data, concluding that nigricapillus and destructus are better treated as two separate biological and recognition species. For more detailed explanations and discussions stemming from historical taxonomic references, please refer to the publication. What follows is a blend of selected copied and reorganized text and images from Areta & Benítez Saldívar (2025) with some minor adjustments to facilitate reading.

 

Songs: After discarding duplicates, Areta & Benítez Saldívar (2025) compiled a total of 57 songs of nigricapillus and 129 songs of destructus that were assessed aurally and through examination of spectrograms. The song of nigricapillus is a rapid series of around 25 clear, pulsated whistled notes that begins with a few more spaced notes that become mostly evenly paced with a slight rise in pitch halfway through the song and a relatively monotonous ending (sigmoid-like spectrographic contour; Fig. 1). The song of destructus is a very rapid eerie series of around 40 ventriloquial notes that fall and rise in pitch and decrease markedly in pace in the second half (smile-like spectrographic contour; Fig. 1). The vocalizations of nigricapillus and destructus have been described accurately in field guides, but the importance of their differences has not been fully realized. The song of nigricapillus was described as "a rapid, pulsating series of ca. 20 deep, resonant, whistled notes, the first 2-3 slower, more staccato, the next 6-8 rising in pitch, the last 10-12 on the same pitch, with the final 2 notes slower, the entire series lasting 4-5 sec" (Stiles and Skutch 1989, p. 287), while that of destructus "resembles that of Rufous-capped Antthrush, but shorter, an eerie, quavering glissando of about 30 notes in 3 sec, sliding upscale and slowing noticeably at the end; ventriloquial" (Hilty and Brown 1986, p. 417). The song of nigricapillus has been aptly likened to the shorter song of Thicket Antpitta Myrmothera dives ( Garrigues and Dean 2007; Vallely and Dyer 2018), a comparison that does not apply to the song of destructus.

 

The quantitative acoustic characterization evidenced that songs of both taxa are 100% diagnosable (n=21 nigricapillus, n=38 destructus). Acoustic data showed that 13 out of 15 variables differed significantly between nigricapillus and destructus, the differences in 10 of these 13 variables were very marked with non-overlapping mean ± SD, automatically indicating that they belong to statistically different populations. Taxon nigricapillus showed lower song peak frequency, longer mean note duration and mean interval between notes, fewer notes per song, slower pace, relatively even pace in the first and second halves of the song, lower peak frequencies of first, median, and final note, and longer duration of first, median, and final note. In contrast, destructus had higher song peak frequency, shorter mean note duration and mean interval between notes, more notes per song, faster pace, a marked deceleration in the second half of the song, higher peak frequencies of first, median, and final note, and shorter duration of first, median, and final note (Table 1). The songs of both taxa were clearly separated in multivariate space (Fig. 2A) and a cluster analysis also revealed two distinctive groups, unambiguously including all destructus samples clustered separately from all nigricapillus recordings (Fig. 2B).

 

The song of nigricapillus remains essentially identical through ca. 490 km across Costa Rica and into C Panama in our sample. t]The song of destructus also remains basically the same through its distribution, ca. 1130 km extending from NC Colombia to SW Ecuador. The diagnostic song types are separated by a gap of ca. 190 km between NW Colombia (Jardín Botánico del Pacífico, Chocó) and E Panamá (Nusagandi, Guna Yala), and exhibit no sign of intermediacy closer to their respective limits (Fig. 1). Further documented records are needed to properly understand the actual distributional gap between nigricapillus and destructus. The width of the gap is at least 190 km as shown by our song-recordings dataset, but possibly much smaller: birds seen, heard, and tape-recorded at Cuchilla del Lago on the Colombian side of the Serranía de Darién in the Cerro Tacarcuna (Renjifo et al. 2017) most likely represent the taxon nigricapillus. This would reduce the gap between nigricapillus (Cuchilla del Lago; see question mark in Figure 1) and destructus (Reserva La Bonga) to 100 km within Colombia. Unfortunately, the sound recordings were not available at the time of our writing and could not be assessed (J. Avendaño in litt.). Further fieldwork should clarify the extent of their allopatry and assess whether the Río Atrato and its formidable swamps act as a biogeographic barrier for these taxa (Haffer 1970, 1975, Renjifo et al. 2017). The Cerro Tacarcuna records, if confirmed to be nigricapillus (which seems very likely), would indicate that both taxa (nigricapillus and destructus) occur in South America.

 

Fig. 1. Plumage aspect, songs, and geographic distribution of songs of Black-capped Antthrush (Formicarius nigricapillus), and Black-hooded Antthrush (F. destructus) analysed in this study. Orange circles: F. nigricapillus (photo: ML-452131711, San Gerardo Biological Station, Costa Rica, by Mark Hebblewhite; song: ML-220516, Reserva Biológica Bosque Nuboso Monteverde, Costa Rica by D. L. Ross). Blue circles: F. destructus (photo: ML-121617911, Rio Silanche Bird Sanctuary, Ecuador by Nick Athanas; song: JIA-10, Reserva de Bosque Seco Lalo Loor, Ecuador by J. I. Areta). Circles with a central dot denote songs measured quantitatively; plain circles denote songs studied aurally (see Appendices 1 and S1); question mark (?) indicates unconfirmed sound recording of nigricapillus from Cerro Tacarcuna in Colombia. The differences in plumage (chestnut nape in nigricapillus, black nape in destructus), song, and morphometrics collectively support the recognition of nigricapillus and destructus as separate species.

 

Pajarografo Sólido:NACHO:Enviados:5-Formicarius destructus y F. hoffmanni:0-Formicarius destructus enviado:0-Reviewed version 15 Nov 2024+20 Jan 2025:Figures corrected for submission after review (chequear):Figure_1 22 Nov 2024 Cuchilla del Lago uncertain 1.jpg

 

 

Table 1. Acoustic parameters of songs of Black-capped Antthrush (Formicarius nigricapillus), and Black-hooded Antthrush (F. destructus). Values shown are mean ± SD [range], n= sample size. Asterisk denotes significant statistical differences in the Mann-Whitney non-parametric test (α<0.05), and plus symbol denotes non overlapping mean ± SD. See Appendix 1 for measured sound recordings.

 

Variable

F. nigricapillus (n=21)

F. destructus (n=38)

Bandwidth 90% (Hz)

205.35±56.37 [93.8-281.2]

207.93±38.45 [140.6-281.2]

Duration 90% (s)*

2.38±0.33 [1.74-3.00]

2.64±0.44 [1.92-3.88]

Peak frequency (Hz)* +

1484.37±61.75 [1406.2-1593.8]

1689.91±61.7 [1546.9-1781.2]

Mean note duration (s)* +

0.06±0.01 [0.05-0.09]

0.04±0.01 [0.02-0.07]

Mean interval between notes (s)* +

0.08±0.01 [0.05-0.10]

0.04±0.01 [0.03-0.06]

Number of notes*+

24.95±1.69 [22-29]

37.62±5.57 [28-52]

Sound density

0.56±0.06 [0.46-0.67]

0.6±0.08 [0.47 - 0.79]

Pace*+

5.98±0.42 [5.44-6.96]

9.61±1.09 [7.76-11.81]

Pace change (second/first half)* +

1.04±0.09 [0.88-1.19]

0.76±0.07 [0.63-0.93]

(8.63±1.2/11.37± 1.08)

[5.61-6.67/5.25-7.58]

(6.18±0.32/5.99±0.66)

[6.82-11.46/9.02-12.82]

Peak frequency first note (Hz)* +

1345.97±68.13 [1218.8-1500]

1650.24±93.23 [1406.2-1781.2]

Peak frequency median note (Hz)* +

1412.91±47.54 [1312.5-1500]

1617.82±58.76 [1453.1-1781.2]

Peak frequency final note (Hz)* +

1477.67±60.46 [1406.2-1593.8]

1703.12±69.11 [1546.9-1875]

Duration first note (s)*

0.04±0.02 [0.01-0.06]

0.02±0.01 [0.01-0.03]

Duration median note (s)* +

0.09±0.02 [0.04-0.14]

0.12±0.03 [0.04-0.10]

Duration final note (s)*

0.14±0.04 [0.10-0.26]

0.11±0.03 [0.06-0.21]

*=non-parametric Mann-Whitney test P-value <.05

+=non-overlapping mean ± SD

 

Plumage: To characterize the external appearance of nigricapillus and destructus, Areta & Benítez Saldívar (2025) examined photographs of 37 museum specimens, including the holotypes of both taxa, and vetted 40 good-quality photographs on the citizen science platforms eBird (ebird.org) and iNaturalist (inaturalist.org). They found that nigricapillus exhibits chestnut-brown hindneck (sometimes extending to neck sides in a handkerchief or semicollar; Figure 1), and typically more chestnut-brown back (Fig. 1), whereas destructus exhibits all-black hindneck and neck sides (Fig. 1), and typically browner back. There is some variation in back colour of specimens (but not in the back-neck contrast that distinguishes taxa), with some nigricapillus being seemingly identical in colour to typical destructus (Chapman 1917, USNM specimens). In some nigricapillus individuals, the chestnut-brown hindneck is extensive and expands onto the sides of the neck creating a semicollar, which gives these individuals a capped aspect, that is less prominent in birds with less extensive chestnut-brown hindneck. On the other hand, all individuals of destructus show a hooded aspect, caused by its wholly black head, hind neck and neck sides (Fig. 1).

 

 

Fig. 2. Quantitative analyses of songs of Black-capped Antthrush (Formicarius nigricapillus), and Black-hooded Antthrush (F. destructus). (A) Plot of the first two principal components (PC1 vs. PC2) of the Principal Component Analysis. Ellipses depict 95% confidence intervals. (B) Dendrogram from the agglomerative hierarchical cluster analysis (UPGMA). Both methods consistently show that nigricapillus and destructus differ markedly in songs supporting their treatment as separate species. See Appendix 1 and S1 for songs measured.

 

Pajarografo Sólido:NACHO:Enviados:5-Formicarius destructus y F. hoffmanni:0-Formicarius destructus enviado:0-Reviewed version 15 Nov 2024+20 Jan 2025:Figures corrected for submission after review (chequear):Figure_2 28_Feb_2025.tiff

 

Morphometry: based on a limited dataset (n=6 nigricapillus, n=9 destructus), Areta & Benítez Saldívar (2025) concluded that 1) bill length (exposed culmen) showed no overlap between taxa, with all individuals of nigricapillus having a longer bill than destructus, and therefore no overlap in mean ± SD values (Fig. 3a), 2) nigricapillus was longer winged than destructus, with exact overlap only in their extreme values, and no overlap in mean ± SD values (Fig. 3b), and 3) tail length was longer in nigricapillus than in destructus, but the difference was not statistically significant (two tailed t-test p=0.17) (Fig. 3c).

 

Fig. 3. Morphological measurements of Black-capped Antthrush (Formicarius nigricapillus), and Black-hooded Antthrush (F. destructus). Figure depicts median and quartiles on box plots. Asterisk denotes significant statistical differences in one-way ANOVA (α<0.05). The morphological differences in bill and wing length coupled to plumage differences give support to the treatment of nigricapillus and destructus as separate species. See Table 2 for sample sizes and data, and Appendix 2 and S2 for specimens measured and studied.

 

Pajarografo Sólido:NACHO:Enviados:5-Formicarius destructus y F. hoffmanni:0-Formicarius destructus enviado:0-Reviewed version 15 Nov 2024+20 Jan 2025:Figures corrected for submission after review (chequear):Figure_3 22 Nov 2024.tiff

 

 

The differences in songs and plumage herein described can be readily appreciated in videos of free-ranging singing birds:

 

nigricapillus: https://macaulaylibrary.org/asset/608419951

destructus: https://macaulaylibrary.org/asset/316228641

 

Taxonomic assessment

The marked differences in vocalizations and morphology, and moderate but consistent plumage differences strongly supports the elevation of the taxon destructus to species-level, leading to the recognition of two allopatric and monotypic species, F. nigricapillus and F. destructus. Areta & Benítez Saldivar (2025) based their taxonomic conclusions on the recognition concept of species (Paterson 1985), whereas the same species would be recognized by applying the biological species concept (Mayr 1963; “isolation concept” fide Paterson 1985). The inferred level of discontinuity between nigricapillus and destructus is of such a magnitude that presumably any other modern species concept would recognise them as separate species, whether based on mating or other important attributes, phenotypic distinctiveness, presence of autapomorphies, or phylogenetic independence (Cracraft 1983, Mishler & Brandon 1987, Gill 2014, Areta et al. 2019, Winker 2021). Howell & Dyer (2022:27) wrote that "Differences in plumage and song indicate that Central American nigricapillus and South American destructus (Choco Antthrush) are best treated as separate species." A view that is amply supported by Areta & Benítez Saldivar (2025).

 

In terms of plumage, the differences between nigricapillus and destructus (Fig. 1) would be among the least conspicuous for two Formicarius species-level taxa, and comparable to (although less obvious than) those between F. moniliger and F. analis. They differ most notably by the presence of a rufous-chestnut fore-collar below the black throat in moniliger, whereas the black throat contacts the grey chest directly in the two subspecies groups of F. analis (Howell 1994, Vallely and Dyer 2018). However, less obvious plumage differences exist between the analis and hoffmanni subspecies groups within F. analis despite their noticeable vocal differences (Howell 1994) which are compatible with species-level differences in the genus (Krabbe and Schulenberg 2003, van Dort et al. 2023, Benítez Saldívar & Areta in prep.).

 

Common names

Most species in the genus Formicarius carry common English names that refer to plumage features. We propose to adopt the name Black-capped Antthrush for F. nigricapillus and Black-hooded Antthrush for F. destructus, which focus on one of the main plumage differences between them and retain a connection to the former Black-headed Antthrush used for the composite species. We find the proposed use of Black-hooded Antthrush for F. nigricapillus by Howell and Dyer (2022) to be misleading, as this taxon is capped rather than hooded. The smaller bill of destructus is difficult to appreciate in field conditions, and bill features do not seem useful to coin common names here. Finally, Choco Antthrush has been proposed for destructus (Howell and Dyer 2022); although a good name, there are other antthrushes in the Choco, and it loses the connection to the former Black-headed Antthrush name.

 

Voting and recommendations

Perhaps the best thing to do is to vote on three different aspects of the proposal:

 

A) Species limits: We recommend a YES vote to split F. destructus from F. nigricapillus.

 

B) English names: We recommend a YES vote to adopt the name Black-capped Antthrush for F. nigricapillus and Black-hooded Antthrush for F. destructus

 

C) Presence of nigricapillus in Colombia. A lingering question is whether the Cerro Tacarcuna records in Colombia should be provisionally considered to be nigricapillus until shown otherwise. For biogeographic reasons, we lean towards the maintenance of nigricapillus in the SACC list based on these records, as it seems unlikely for destructus to reach the Cerro Tacarcuna crossing the Atrato swamps. We recommend a YES vote to keep F. nigricapillus in the SACC list.

 

References

Areta JI, Depino EA, Salvador SA, Cardiff SW, Epperly K, Holzmann I (2019) Species limits and biogeography of Rhynchospiza sparrows. Journal of Ornithology 160:973-991

Areta, J.I., Benítez Saldívar, M.J. (2025). Species limits in the Black-headed Antthrush (Formicarius nigricapillus). J Ornithol https://doi.org/10.1007/s10336-025-02265-5

Chapman FM (1917) The Distribution of Bird-Life in Colombia. A Contribution to a Biological Survey of. South America. Bull. Amer. Mus. Nat. Hist., 36.

Cory CB, Hellmayr CE (1924) Catalogue of Birds of the Americas. Publications of the Field Museum of Natural History (Zoological Series). Volume 13, Part 3. Chicago.

Cracraft J (1983) Species concepts and speciation analysis. Curr Ornithol 1:159–187

Garrigues R, Dean R (2007) The Birds of Costa Rica. Zona Tropical, Comstock Publishing Associates, Cornell University Press.

Gill F (2014) Species taxonomy of birds: which null hypothesis? Ornithological Advances 131: 150–161

Haffer J (1970) Geologic-climatic history and zoogeographic significance of the Urabá region in northwestern Colombia. Caldasia 10: 603–636.

Haffer J (1975) Avifauna of northwestern Colombia, South America. Bonner Zoologische Monographien 7: 1–182.

Hartert E (1898) On a collection of birds from north-western Ecuador, collected by Mr. W. F. H. Rosenberg. Novitates Zoologicae 5: 477505.

Hartert E (1902) Some further notes on the birds of north-west Ecuador. Novitates Zoologicae 9: 599617.

Hilty SL, Brown WL (1986) A Guide to the Birds of Colombia. Princeton: Princeton University Press.

Howell SNG (1994) The specific status of black-faced antthrushes in Middle America. Cotinga 1: 2025.

Howell SNG, Dyer D (2022). Costa Rica even richer than we thought? North American Birds 73: 22–33.

Krabbe NK, Schulenberg TS (2003). Black-headed Antthrush (Formicarius nigricapillus). In Handbook of the Birds of the World Alive (del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E, eds). Lynx Edicions, Barcelona.

Krabbe NK, Schulenberg TS (2020). Black-headed Antthrush (Formicarius nigricapillus), version 1.0. In Birds of the World (del Hoyo J, Elliott A, Sargatal J, Christie DA, de Juana E, eds). Lynx Edicions, Barcelona. Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.blhant1.01

Mayr E (1963) Animal species and evolution. Harvard University Press, Cambridge

Paterson HEH (1985) The recognition concept of species. In: Vrba ES (ed) Species and speciation. Transvaal Museum Monograph, Pretoria.

Mishler BD, Brandon RN (1987) Individuality, pluralism, and the Phylogenetic Species Concept. Biology and Philosophy 2: 397414

Renjifo LM, Repizo A, Ruiz-Ovalle JM, Ocampo S, Avendaño JE (2017) New bird distributional data from Cerro Tacarcuna, with implications of conservation in the Darién highlands of Colombia. Bulletin of the British Ornithologists´ Club 137: 4666.

Ridgway R (1893) A revision of the genus Formicarius Boddaert. Proceedings of the United States National Museum 16: 667686.

Ridgway R (1911) The Birds of North and Middle America: A Descriptive Catalogue. Part V. Bulletin of the United States National Museum 50.

Salvadori T, Festa E. 1899. Viaggio del Dr. Enrico Festa nell’Ecuador. XXI. Uccelli. Parte seconda - Passeres clamatores. Bollettino dei Musei di Zoologia ed Anatomia Comparata della R. Università di Torino 15 (362): 134.

Stiles FG, Skutch AF (1989) A Guide to the Birds of Costa Rica. Ithaca, NY: Cornell University Press.

Vallely AC, Dyer D (2018) Birds of Central America. Princeton University Press, Princeton and Oxford.

van Dort J, Patten MA, Boesman PFD (2023). Black-faced Antthrush (Formicarius analis), version 2.0. In Birds of the World (S. M. Billerman, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bow.blfant1.02

Wetmore A (1972). The Birds of the Republic of Panamá. Volume 3. Passeriformes: Dendrocolaptidae (Woodcreepers) to Oxyruncidae (Sharpbills). Smithsonian Miscellaneous Collections 150(3). Smithsonian Institution, Washington, D.C., USA.

Winker K (2021) An overview of speciation and species limits in birds. Ornithology 138: 127 10.1093/ornithology/ukab006

 

Links to supplementary information:

https://static-content.springer.com/esm/art%3A10.1007%2Fs10336-025-02265-5/MediaObjects/10336_2025_2265_MOESM1_ESM.xlsx

 

https://static-content.springer.com/esm/art%3A10.1007%2Fs10336-025-02265-5/MediaObjects/10336_2025_2265_MOESM2_ESM.xlsx

 

Addendum:

Genetics: The coincidental break in plumage and vocalizations between F. moniliger and F. analis umbrosus (a representative of the hoffmanni group) were used as arguments to support the elevation of moniliger to species status (Howell 1994). More recently, phylogenetic data have shown that F. moniliger is sister to a clade including F. destructus as more closely related to F. analis (including representatives of the hoffmanni and analis subspecies groups) (Harvey et al. 2020). This distant relationship between moniliger and analis further reinforces the species-level split of F. moniliger and lends support to vocal differences as a useful tool to establish species limits in Formicarius. Although there are no available genetic data for nominotypic nigricapillus, the vocal distinctions between nigricapillus and destructus seem as or more marked than those between F. moniliger and F. analis of the analis and hoffmanni subspecies groups. In our paper we predicted that nigricapillus and destructus will exhibit levels of genetic differentiation tantamount to their vocal and morphological distinctions.

 

The Formicarius phylogenetic tree from Harvey et al. 2020:

 

Pajarografo Sólido:Users:javierareta:Desktop:Screen Shot 2025-04-04 at 3.48.51 PM.png

 

Harvey MG, Bravo GA, Claramunt S, et al. (2020) The evolution of a tropical biodiversity hotspot. Science 370: 1343–1348.

 

 

Juan I. Areta and M. Juliana Benítez Saldívar, April 2025

 

 

 

Voting chart: https://www.museum.lsu.edu/~Remsen/SACCPropChart1044+.htm

 

 

Comments from Remsen:

“A. YES. Although playback trials would be nice, the degree of difference in the songs is consistent with species rank, and burden-of-proof falls on those who would treat them as subspecies.

“B. YES, for all the good reasons outlined in the proposal, especially maintaining the connection between old and new names.  Also, follows SACC guidelines (https://www.museum.lsu.edu/~Remsen/SACCEnglishNameGuidelines.pdf) for providing new names for both daughters in a parental split.  After having looked through Macaulay photos of dozens of destructus, “hooded” seems especially appropriate for the extent of the black.  However, as for nigricapillus, a problem is that “capped” is typically used for cases in which the bird has a much more distinctive color patch on the top of the head, which is not really the case for nominate ruficapillus, despite the scientific name.  (TO BE MODIFIED)

“C. YES, but somewhat reluctantly because the recordings are unavailable and because perhaps the Colombian national committee should have the say on this one.  On the other hand, to propose that the records are not nigricapillus makes no biogeographic sense, and I agree with this statement in the proposal “Cerro Tacarcuna records in Colombia should be provisionally considered to be nigricapillus until shown otherwise.”

 

“Anecdote: puzzled by the name destructus, I looked at Jobling (online), and here is the derivation from Hartert’s 1898 description: ‘very much destroyed by the shot, but still quite recognizable, is so considerably smaller and darker brown above than typical analis that I do not hesitate to distinguish it subspecifically’.”

 

Comments from Lane: “I must admit this one was not on my radar (although I have seen, and heard, both taxa). The voices are much more distinct that I was expecting, and I think the results of the study presented are good.

 

“A) YES to splitting the two members of the complex as biological species.

“B) The two daughter species are not all that strongly differentiated by plumage, and really can't see F. nigricapillus having a distinctive black crown set off against the rest of the head (I suspect the name was coined because it was blacker-crowned than F. analis... and granted, I am looking at the poorly-lit Macaulay photos rather than specimens to make this statement), but unless we want to get derailed by another English name situation, I will go with the names suggested: Black-crowned and Black-hooded.

”C) YES to retaining both species in the SACC area due to the records from the border near Panama.”