Research Article |
Corresponding author: Luiz A. Rocha ( lrocha@calacademy.org ) Academic editor: Nina Bogutskaya
© 2017 Luiz A. Rocha, Hudson T. Pinheiro, Matt Wandell, Claudia R. Rocha, Bart Shepherd.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rocha L, Pinheiro H, Wandell M, Rocha C, Shepherd B (2017) Roa rumsfeldi, a new butterflyfish (Teleostei: Chaetodontidae) from mesophotic coral ecosystems of the Philippines. ZooKeys 709: 127-134. https://doi.org/10.3897/zookeys.709.20404
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A new species of the butterflyfish genus Roa is herein described from the Verde Island Passage in the Philippines, West Pacific Ocean. Roa rumsfeldi sp. n. was found on mesophotic coral ecosystems at Puerto Galera and Batangas, and sampled through technical mixed-gas rebreather diving at 100–130 m depth. This represents the fifth known species of Roa. The main differences between Roa rumsfeldi sp. n. and its congeners are the lower number of pored scales in the lateral line, longer snout, longer caudal peduncle, shorter caudal fin, pelvic fin color (dark first spine vs. white in all other Roa), and genetics (8.4% divergence from its closest relative Roa modesta in the mitochondrial COI gene). Roa spp. are usually seen in pairs, but the two specimens we collected were solitary individuals. We have kept one of the specimens alive in the California Academy of Sciences’ Twilight Zone exhibit for more than one year, where it thrives and is feeding on a variety of dried and fresh food.
Coral triangle, deep reefs, new species, rebreather diving, reef fish
While collecting live specimens in Anilao, Philippines to display at the California Academy of Sciences’ Twilight Zone: Deep Reefs Revealed exhibit in 2016, LAR collected a butterflyfish identified at the time as Roa cf. modesta. After the specimen arrived in San Francisco and entered quarantine, MW noticed that the recent arrival differed from our previously collected specimens by having a black spine in the ventral fin (white in others). We went through our previous years’ collections and found a second specimen of this species collected by spear in 2015 in Puerto Galera, Philippines.
These specimens represent an undescribed species of the genus Roa Jordan, 1923, which currently contains four species, all from mesophotic coral ecosystems (MCEs; 30-150m depth) in the Indo-Pacific. Although some taxonomists consider this genus as a “modestus species complex” within Chaetodon (
The currently recognized species of Roa have largely allopatric distributions: Roa modesta (Temminck & Schlegel, 1844) was described from Japan and is distributed along the northwestern Pacific, recorded in the Philippines, Hong Kong, Taiwan, Ryukyu and Ogasawara Islands, and Nagasaki, in Japan (
The specimens were collected using hand nets or a Hawaiian sling while diving on mixed-gas, closed-circuit rebreather (Hollis Prism 2). Counts were performed using a microscope, and morphological characters were measured to the nearest 0.1 mm following
Proportional measurements of Roa species. Values that do not overlap between Roa rumsfeldi sp.n. and the other species are in bold.
Roa rumsfeldi sp. n. | Roa australis | Roa excelsa | Roa modesta | Roa jayakari | ||
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Standard length | 64.2 mm | 69.5–119 mm | 94–105 mm | 51.5–97 mm | 99–104.3 mm | |
% | % | % | % | % | ||
Body | depth in SL | 69.8 | 63.6–75.0 | 61.0–67.0 | 66.5–73.2 | 72.5–77.5 |
width in SL | 15.4 | 14.0–19.4 | 14.9–17.3 | 15.5–19.3 | 15.6–16.5 | |
Head | length in SL | 41.1 | 36.1–42.0 | 34.9–37.1 | 31-5–40.9 | 40.2–40.9 |
Snout | length in HL | 35.9 | 28.2–34.6 | 29.3–34.5 | 32.5–32.7 | 33.5–35.5 |
Eye | diameter in HL | 32.2 | 30.1–36.1 | 33.6–32.5 | 30.8–32.6 | 23.6–27.1 |
Interorbital | width in HL | 27.5 | 19.6–27.3 | 21-7–25.3 | 22.7–27.8 | 22.7–23.9 |
Caudal peduncle | depth in SL | 10.7 | 11.3–14.1 | 10.2–11.1 | 12.0–13.3 | 10.8–11.2 |
length in SL | 8.46 | 4.6–6.4 | 4.2–4.9 | 4.6–5.5 | 6–8.2 | |
Caudal fin | length in SL | 15.9 | 21-8–25.9 | 20.0–21.2 | 21.1–25.6 | 19.8–22.3 |
Pectoral fin | length in SL | 29.1 | 29.4–36.2 | 28.8–33.6 | 26.3–30.2 | 27.6–28.3 |
Dorsal-fin base length | spinous in SL | 40.0 | 35.8–42.0 | 36.4–40.1 | 36.2–38.8 | 38.8–42.6 |
Dorsal-fin base length | soft in SL | 31.1 | 31.7–42.0 | 31.1–33.9 | 35.3–40.8 | 33.4–33.6 |
Dorsal-fin spine length | 1st in SL | 9.1 | 6.2–8.8 | 7.0–7.7 | 6.4–9.3 | 8.4–9 |
2nd in SL | 18.3 | 13.4–21.0 | 15.0–17.3 | 15.9–22.3 | 16.6–18.3 | |
3rd in SL | 24.5 | 35.4–21.3 | 33.9–36.5 | 23.4–30.6 | 24.9–25 | |
4th in SL | 29.1 | 37.1–25.6 | 26.5–32.6 | 26.3–33.7 | 26–28.8 | |
Dorsal-fin soft-ray length | 1st in SL | 12.0 | 17.6–23.9 | 16.6–21.2 | 19.5–23.4 | 23.2–26.3 |
Anal-fin base | length in SL | 34.2 | 31.2–37.4 | 30.6–32.6 | 36.0–41.8 | 35.1–36.1 |
Anal-fin spine length | 1st in SL | 12.9 | 9.3–13.7 | 12.1–13.2 | 9.7–11.9 | 13.4–15.3 |
2nd in SL | 25.2 | 21.8–28.1 | 27.1–30.0 | 18.7–19.6 | 21.3–24.3 | |
3th in SL | 12.8 | 18.9–23.0 | 17.2–20.8 | 19.6–21.9 | 16.7–20.4 | |
Anal-fin soft-ray | longest in SL | 13.8 | 17.5–23.2 | 18.5–22.3 | 17.5–26.2 | 21.4–23 |
Ventral fin | length in SL | 27.5 | 27.5–38.1 | 29.9–33.0 | 30.2–37.8 | 26.2–30.8 |
spine length in SL | 25.2 | 20.3–30.2 | 23.6–26.0 | 22.3–25.8 | 23.3–27.7 |
Puerto Galera, Philippines.
Roa modesta
Roa rumsfeldi sp. n. differs from all of its congeners by the smaller number of pored scales in the lateral line pored scales (27 versus 37-46 in other Roa), longer snout (35.9% in HL vs. 28.2–35.5%), shorter first dorsal ray (12% in SL vs. 16.6–26.6%), longer caudal peduncle (8.5% in SL vs. 4.2–8.2%), shorter caudal fin (15.9% in SL vs. 19.8–25.9%), and a dark brown pelvic-fin spine (white in all other known Roa species).
Dorsal fin rays XI, 20, last soft ray branched to the base and counted as one; dorsal-fin base length: spines 40.0% in SL, and soft rays 31.1% in SL; spines increase in height from the first to the fourth: length of first spine 9.1% in SL, length of second spine 18.3% in SL, length of third spine 24.5% in SL, length of fourth spine 29.1% in SL. Soft dorsal fin follows sharp descent of rear portion of body. First soft ray 12.0% in SL. Anal-fin rays III, 17, last soft ray branched to the base and counted as one; anal-fin base 34.2% in SL; second spine very long, 25.3% in SL. Pectoral fin 13; length 29.1% in SL (Table
Origin of dorsal fin above the origin of pectoral fin, posterior end of head; fin base long and mostly horizontal; soft part of dorsal fin curves downward from the third ray to caudal peduncle; soft rays posterior margin vertical; origin of anal fin below 9th spine of dorsal fin; pelvic fin with strong spine and filamentous first soft ray.
Body and head with ctenoid scales, becoming smaller towards nape and snout. Scales extending to about one third of the median fins. Tubed scales of lateral line rising at a steep angle from origin with 22 scales in an almost straight line, bending abruptly downward, ending behind posterior third of soft dorsal fin.
Roa rumsfeldi sp. n. (Figure
As described above in color in life, but with lighter brown bands.
We name Roa rumsfeldi to honor Donald Rumsfeld who immortalized the quote: “there are known knowns; there are things we know we know. We also know there are known unknowns; that is to say we know there are some things we do not know. But there are also unknown unknowns – the ones we don’t know we don’t know.” He said that when referring to the uncertainties of war, but we think it applies perfectly to the taxonomy of MCE species: We only realized this species was new after we took a good look at it here at the aquarium in San Francisco, so we think it’s a perfect example of an unknown unknown.
Roa rumsfeldi sp. n. is only known to occur in the Verde Island Passage, central Philippines. It was found on MCEs of Puerto Galera, Oriental Mindoro, and Bauan, Batangas, between 100 and 130 m depth. However, the species likely has a wider distribution and remains undetected because of its preferred depth range. The ecosystems where it was found vary from sheltered rocky outcroppings heavily covered by fine sediment to areas exposed to strong currents. The ambient seawater temperature varied between 19 and 21°C during our dives, which were conducted in April-May over several years. Azooxanthellate gorgonians, black corals, and solitary stony corals are the most abundant benthic invertebrates in this habitat.
In addition to the singular characteristics presented in the diagnosis section, Roa rumsfeldi sp. n. differs from its congeners by a lower number of dorsal soft rays (20) than R. jayakari (22–24); longer head (41.0% in SL) than R. excelsa (34.9–37.1); fewer dorsal-fin rays (20) and shorter dorsal-fin rays base (31.1% in SL) than R. modesta (22 rays and 35.3–40.8% in SL); and shorter 3rd anal-fin spine (12.8% in SL) and longest anal-fin ray (13.8% in SL) than R. australis (17.6–23.9, 18.9–23.0 and 17.5–23.2% in SL, respectively). Moreover, the COI gene sequence of Roa rumsfeldi does not match any other Roa species available at Genbank. The uncorrected genetic divergence at the COI gene between R. rumsfeldi and the two other available Roa (R. jayakari and R. modesta) is 10.5% and 8.4%, respectively.
Despite a few records of Roa modesta and R. excelsa in shallow waters of Japan and Hawaii, Roa species are normally found at depths exceeding 100 m, with trawling being the most common sampling method for these species (
This work was funded by the generous support of donors who endorsed the California Academy of Sciences’ Hope for Reefs Initiative and through a grant from the National Science Foundation to T. Gosliner, R. Mooi, G. Williams and L.A. Rocha (DEB 12576304). We are grateful to many colleagues who helped in the field, lab, and with discussions: M. Bell, B. Greene, E. Jessup, M. Lane, N. Nazarian, T. Phelps, R. Pyle, and G. Rocha. Hollis and Anilao Beach Club provided gear and logistical support. The Philippines research and collecting permits (GP-0085-15 and GP-0112-16) were provided by the Bureau of Fisheries and Aquatic Sciences. This is a collaborative research initiative with key Philippine partners including: former Secretary of Agriculture P. J. Alcala; former Philippine Consul General M. Paynor and the Consular staff in San Francisco; former BFAR Director A. G. Perez; BFAR colleagues, especially A. Vitug and L. Labe; and NFRDI colleagues especially, Acting Director D. Bayate and N. Romena.