diff --git a/data/6A/DC/2A/6ADC2AFAB9DC5DCDB5B1CEF1B61F1C4E.xml b/data/6A/DC/2A/6ADC2AFAB9DC5DCDB5B1CEF1B61F1C4E.xml new file mode 100644 index 00000000000..19f1abd5d53 --- /dev/null +++ b/data/6A/DC/2A/6ADC2AFAB9DC5DCDB5B1CEF1B61F1C4E.xml @@ -0,0 +1,1041 @@ + + + +Description of a new species of Stauropathes (Anthozoa, Antipatharia, Schizopathidae) from Puerto Rico + + + +Author + +Horowitz, Jeremy +0000-0002-2643-5200 +Department of Invertebrate Zoology, Smithsonian Institution, National Museum of Natural History, Washington, DC, USA + + + +Author + +Barajas, Mina +Department of Zoology, California State Polytechnic University, Humboldt, Arcata, CA, USA + + + +Author + +McCartin, Luke J. +0000-0001-5374-3148 +Department of Biological Sciences, Lehigh University, Bethlehem, PA, USA + + + +Author + +Vohsen, Samuel A. +0000-0003-1710-292X +Department of Biological Sciences, Lehigh University, Bethlehem, PA, USA + + + +Author + +Herrera, Santiago +0000-0001-7204-9434 +Department of Biological Sciences, Lehigh University, Bethlehem, PA, USA + +text + + +ZooKeys + + +2025 + +2025-03-13 + + +1231 + + +331 +346 + + + +journal article +10.3897/zookeys.1231.136967 +4ECF017A-A679-4DAE-89BE-6606835C0F44 + + + + + +Stauropathes monopinnata +Horowitz & Barajas + +sp. nov. + + + + +Figs 1 +, +3 +, +4 +, +5 +, +6 +, +Table 1 +, Suppl. material 1 + + + + +Material examined. + + + + +Holotype + +• +USNM 1689279 +, +Whiting Bank +, +Puerto Rico +, + +17.8398°N +, +65.6976°W + +, + +738 m +depth + +. Schmidt Ocean Institute R/V +Falkor (too) +, FKt 230417, Health diagnostic of deep-sea coral, ROV +SuBastian +dive 518, + +April 29, 2023 + +( +SEM +stub No. 538) + +. + + +Paratype + +• +USNM 1424060 +, +250 km southeast of +Midway Atoll +, +Hawaii +, + +26.65°N +, +175.4°W + +, + +1604 m +depth + +. NOAA (National Oceanic and Atmospheric Administration) R/V +Okeanos Explorer +, Cruise EX 1603, Hohonu Moana: Exploring Deep Waters off Hawaii, ROV +Deep Discoverer +dive 5, + +March 5, 2016 + +( +SEM +stub 539) + +. + + + + +Type locality. + + +Whiting Bank, +Puerto Rico +, +738 m +depth. + + + + +Diagnosis. + + +Corallum monopodial, planar, unbranched, and pinnulate to the first order. Pinnules in two anterolateral rows and arranged in subopposite (almost opposite) pairs. Lowermost pinnules on stem +8 cm +or more in length, decreasing in length towards apex of colony. Spines smooth and triangular with a rounded apex, +0.05–0.08 mm +tall. Polyps +4–9 mm +in transverse diameter, +2–3 mm +interpolypar space, four to eight polyps counted per +5 cm +. + + + + + +Description of +holotype +. + + + +The +holotype +( + +USNM +1689279 + +) is a +15 cm +tall monopodial colony (Fig. +3 A – C +). The unpinnulated section is about +2 cm +based on in situ imagery because the basal plate and lowermost section of unpinnulated stem were not collected (Fig. +3 C +). The pinnulated section is +13 cm +in length. Pinnules are simple, arranged in two anterolateral rows and in subopposite pairs (Fig. +3 A, B +). The pinnules are curved towards the abpolypar side of the colony. The lowermost pinnules on the stem are +8.3 cm +in length, pinnules midway up the pinnulated section of the stem are +8 cm +in length, and the uppermost pinnules on the stem are +4.2 cm +in length. Diameter of the lowermost pinnule near the base of the pinnule is ~ +1 mm +. Distance between pinnules in one row ranges from +1.3 to 1.7 cm +and distance between pinnules of the same pair (i. e., on opposite sides of the stem) is < +1 mm +, in some cases nearly opposite (Fig. +3 B +). Eight pinnules can be counted per +5 cm +including pinnules in both rows. The distal angle of pinnules is 80–90 ° except for the uppermost pinnules, which are about 45 ° (Fig. +3 A – C +), and the interior angle formed by the subopposite pinnules is 160–180 ° (Fig. +3 A +). + + + + + + + +Stauropathes monopinnata + +sp. nov. +holotype: +USNM 1689279 +A +collected colony with +B +blue rectangle showing zoomed-in view of pinnule pattern +C +in situ image of colony +D +in situ image of colony showing zoomed-in view of polyp characteristics +E +section of pinnule showing polyp characteristics. + + + +Polyps are arranged in a single row (Fig. +3 C, D +). Polyps are +8–9 mm +in the transverse diameter with +0.2–0.3 mm +interpolypar space, resulting in polypar density ranging from 4 to 5 per +5 cm +(Fig. +3 E +). Based on in situ imagery, tentacles exceed +1 cm +when fully extended and the tissue color is dark red, and when preserved is dark red to dark brown. + + +Spines (Fig. +4 A – C +) are smooth and triangular with a rounded apex (Fig. +4 C +) that is at right angles to the axis or slightly inclined distally, with long and sloping distal and proximal edges. Polypar and abpolypar spines are +0.03 to 0.05 mm +tall. The spines on pinnules are arranged in longitudinal rows, six to seven of which can be seen in one view (Fig. +4 A, B +). Spines are spaced +0.12–0.33 mm +apart in a row, with about four to five spines per mm (Fig. +4 A, B +). + + + + + + + +Stauropathes monopinnata + +sp. nov. +holotype: +USNM 1689279 +A, B +sections of pinnules showing skeletal spines +C +zoomed-in view of a singular skeletal spine. + + + + + + +Description of the +paratype +. + + + +The +paratype +( + +USNM +1424060 + +) is a +14 cm +tall colony (lowermost section of stem not collected) and the pinnulated section of stem is +12 cm +(Fig, 5 A). Based on in situ imagery ( + +https://data.oceannetworks.ca/SeaTube?resourceTypeId=1000&resourceId=23621&diveId=3000&time= +2016-03-06 +T00:56:55.000Z + +), the whole colony was approximated to be ~ +25 cm +in length and the unpinnulated section of stem was ~ +7 cm +in length. Striatum is present and distinct from the lower broken-off end of the stem to the lowermost pair of pinnules. Pinnules are simple, arranged in anterolateral rows and in subopposite pairs (Fig. +5 A +). The lowermost pinnules are 2.5 and 3.0 cm long; however, based on in situ imagery (Fig. +5 B +) the lowermost pinnules were the longest on the colony prior to subsampling. Pinnules midway up the pinnulated section of the stem are 8.0 cm long and the most distal pinnules are +1.3 cm +long (Fig. +5 A +). Diameter near the base of the pinnule is ~ +1 mm +. Distance between pinnules in one row ranges from +1.2 to 1.4 cm +and distance between members of the subopposite pairs is < +1 mm +(Fig. +5 A +). In some cases, the members of a pair appear to be directly opposite to one another. Ten pinnules can be counted per +5 cm +including pinnules in both rows (Fig. +5 A +). Distal angle of pinnules is 80–90 ° and the interior angle formed between pinnules of a subopposite pair is 160–180 ° (Fig. +5 A +). + + + + + + + +Stauropathes monopinnata + +sp. nov. +paratype ( +USNM 1424060 +) +A +collected colony +B +in situ image of colony +C, D +sections of pinnules showing skeletal spines +E +zoomed-in view of a singular skeletal spine. + + + + + + + +Maximum likelihood phylogeny of antipatharians of the families +Schizopathidae +and +Cladopathidae +based on a 50 % complete matrix containing 794 loci. Taxa in green rectangle represent the holotype and paratype specimens of the new species. Ultrafast Bootstrap support values are 100 % at all nodes unless noted with an asterisk (*). Red branches were artificially shortened and do not represent true branch length. The phylogeny rooted to +Cladopathidae +. + + + +Polyps are arranged in a single row and are +4–6 mm +in transverse diameter with an interpolypar space of +2 mm +. Polypar density ranges from seven to eight per +5 cm +. Based on in situ imagery, the tissue is dark red, and when preserved, it is brown to dark brown (Fig. +5 B +). + + +Spines are smooth, laterally compressed, and triangular with a rounded apex (Fig. +5 C – E +). Spines are +0.03–0.06 mm +tall. The spines on pinnules are arranged in longitudinal rows, seven to eight of which can be seen in one view (Fig. +5 C, D +). Spines are spaced +0.23–0.36 mm +apart in a row with about three to four spines per mm (Fig. +5 C, D +). + + + + +Intraspecies variation. + + +Both specimens possess simple pinnules with wide distal angles, arranged in slightly anterolateral rows, and in subopposite pairs that are almost opposite. Spacing between pinnules in one row is wide in both specimens: ≤ +1.7 cm +in the +holotype +and ≤ +1.4 cm +in the +paratype +, resulting in eight and ten pinnules per +5 cm +. The spines on both specimens are short (≤ +0.06 mm +tall), smooth, triangular with a rounded apex, and arranged in longitudinal rows, seven to eight of which are visible in lateral view. Tissue color is similar for both specimens; dark red in situ and a dark red-to-brown when preserved. + + +A minor difference between the +holotype +and +paratype +is that based on in situ footage, the +paratype +pinnules are more rigid than the +holotype +. A major difference between the +holotype +and +paratype +is their polyp characteristics, where the +holotype +has polyps +8–9 mm +in transverse diameter with +0.2–0.3 mm +interpolypar space, resulting in a density of four to five polyps per +5 cm +, while the paratype’s smaller polyps are +4–6 mm +in transverse diameter with +0.2 cm +interpolypar space, resulting in a higher density of seven to eight per +5 cm +. This is a notable difference that could be interpreted as the +holotype +and +paratype +representing different species. However, without additional specimens possessing similar morphological traits as these specimens with differing-sized polyps, it is premature to describe these +two specimens +as polyp size could vary within the species. + + + + +Phylogenetic results. + + +A total of 63–1052 conserved element loci were obtained per specimen. Total number of contigs ranged from 234,560 to 439,048,041 base pairs (bp) (average lengths ranged from 109 to 953 bp). The 50 % taxon-occupancy matrix included 794 loci that were concatenated into an alignment with a total length of 385,232 bp. A 75 % taxon-occupancy matrix, including 465 loci, was also run to compare results, and the topology in the region of the trees, including the new species, did not change. Read and locus summary statistics are detailed in Suppl. material +1 +. + + +The +two specimens +representing the new species with 100 % branch support fell sister to the clade containing + +Stauropathes staurocrada + +( + +USNM +98846 + +species- and genus-level +holotype +specimen, and + +USNM +1071042 + +) (Fig. +5 +), supporting our decision to place the new species in + +Stauropathes + +. This clade also contains + +Stauropathes +cf. +punctata +( +Roule, 1905 +) + +( + +USNM +1606527 + +), + +Stauropathes +cf. +stellata +Opresko, 2019 + +( + +USNM +1424179 + +), and + +Bathypathes alaskensis + +( + +USNM +1013749 + +). + + + + +Comparative diagnosis. + + + +Stauropathes monopinnata + +sp. nov. +differs from the four other species in the genus morphologically. The most prominent difference is that the new species is unbranched whereas the other species are branched. The new species also has wider distances between pinnules in a row; reaching +1.7 cm +in the +holotype +compared to a maximum of +1.2 cm +in + +S. staurocrada + +and + +S. arctica +( +Lütken, 1871 +) + +, and +0.8 cm +in + +S. stellata + +and + +S. punctata + +. Four of the species in the genus possess small spine heights less than +0.07 mm +(spine measurements were not reported for the type of + +S. punctata + +). The species however, differ in the number of rows of spines that can be seen in one lateral view of a pinnule. The number of rows for the new species (six to eight visible on a pinnule diameter of +0.25 mm +) is greater than that for + +S. staurocrada + +(four to six on pinnule diameter of +0.28 mm +) and + +S. stellata + +(three to four on pinnule diameter of +0.28 mm +) and less than + +S. arctica + +(nine to ten on pinnule diameter of +0.34 mm +). The new species also has polyps that are +4 to 9 mm +in transverse diameter, equal to or larger than + +S. staurocrada + +( +4 mm +), and the range includes polyps of + +S. stellata + +( +6 mm +) and + +S. arctica + +( +7 mm +). The transverse diameter of the polyps was not reported for + +S. punctata + +. A complete comparison of the morphological features of + +Stauropathes +species + +can be found in Table +1 +. + + + + + + +Comparison of species in the genus + +Stauropathes + +. + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
Feature + +S. monopinnata + +sp. nov. + + +S. monopinnata + +sp. nov. + + +S. staurocrada + +c + + +S. stellata + +d + + +S. punctata + +e + + +S. arctica + +f +
+holotype +a + +paratype +b +
Corallumunbranchedunbranchedbranchedbranchedbranchedbranched
Stem length (cm) (pinnulated / unpinnulated)13 / 212 / 2 (unpinnulated section incomplete)13 / not collected6.3 / 6.121 / not collected20 / 5
Pinnule diameter near base (mm)1110.51Not reported
Pinnular angle (distal / interior)45–90 ° / 160–180 °80 – 90 ° / 160–180 °60 – 70 ° / 90–150 °80 – 90 ° / 160–180 °80 – 90 ° / 160–180 °80 – 90 ° / 160–180 °
Distance between pinnules on one side (mm)13–1712–148–125–106–812
Max Pinnule length (cm):8.3825.513
Pinnule density per 5 cm (both rows)8105–68Not reportedNot reported
Number of orders of pinnules11111 +> 1? +g +
Spine height (mm)0.03–0.050.03–0.060.04–0.060.06–0.07Not reported0.02–0.06
Spine ornamentationSmoothSmoothSmoothSmoothNot reportedSmooth
Number of spine rows per view6–77–84–63–4Not reported9–10
Space between spines in one row (mm)0.12–0.170.11–0.440.12–0.240.21–0.33Not reported0.06–0.12
Spine density per 1 mm4–5484–6Not reported5
Polyp transverse diameter (mm)8–94–62–45–6Not reported3–7
Polyp density per 5 cm4–57–810–116–7Not reportedNot reported
Number of polyps between adjacent pinnules in the same row222Not reportedNot reportedNot reported
StriatumLowest section not collectedStriations present until first row of pinnulesPresentPresent from 2 cm above basal plate and extends 4 cmNot reportedNot reported
+ + + +a + +Stauropathes monopinnata + +sp. nov. +holotype +( + +USNM +1689279 + +) herein described. +b + +Stauropathes monopinnata + +sp. nov. +paratype +( + +USNM +1424060 + +) herein described. +c + +Stauropathes staurocrada + +holotype +( + +USNM +98846 + +) in +Opresko (2002) +. +d + +Stauropathes stellata + +holotype +( + +USNM +16059 + +) in +Opresko (2019) +. +e + +Stauropathes punctata + +type series described in +Roule (1905) +. +f + +Stauropathes arctica + +Brook’s (1889) +translation of species described by +Lütken (1871) +. +g +Unclear if ramification orders are pinnulated branches or pinnules. + + + +While the lack of branches and the two rows of subopposite pinnules in the new species is typical of + +Bathypathes +species + +, it differs morphologically in several ways: the distances between the members of each subopposite pair are smaller, and in some cases, the pinnules are almost opposite; the colonies have wider spaces between pinnules in a row, and the longest pinnules are also the lowermost ones on the stem. + + + + +Etymology. + +The specific name derives from the Latin “ mono ” (one) and “ pinnata ” (feathered) referring to the new species general appearance due to the distinctive lack of branches compared to the other species in the genus. + + + +Distribution. + + +Known from North Central Atlantic Ocean to North Pacific Ocean from +738 to 1604 m +depth. + + + + +Discussion and conclusions. + + +This study presents morphological and molecular evidence to support the description of a new species within the genus + +Stauropathes + +. Furthermore, this study provides the most speciose molecular phylogeny of the family +Schizopathidae +to date, including specimens representing all 13 accepted schizopathid genera, +holotypes +or +paratypes +of 20 species, five of which also represent types at the genus level, and 12 species sequenced for the first time. + + +The new + +Stauropathes +species + +, which lacks branches (like + +Bathypathes + +), required an emendation of the diagnosis of + +Stauropathes + +to include unbranched morphotypes. Additionally, the finding of one + +Bathypathes +species + +in the + +Stauropathes + +clade and two + +Stauropathes +species + +in the + +Bathypathes + +clade suggests they have a complicated evolutionary history, possibly driven by convergent evolution or hybridization. + + +Speciation is complex, and the phylogenetic models used in black coral studies have relied on maximum likelihood analyses with General Time Reversible substitution model, which does not account for processes like hybridization, recombination, or site-specific variation in substitution rates ( +Steenwyk et al. 2023 +). When dealing with complicated evolutionary histories, which seems to be the case for genera in +Schizopathidae +(see also ‘ The Trigeneric Complex’ described in +Bledsoe-Becerra et al. (2022) +) a multispecies coalescent model should be used for estimating phylogenies while accounting for unresolved lineage sorting ( +Ramírez-Portilla and Quattrini 2023 +). This is an essential next step for resolving relationships at the species level for black corals. + + + + \ No newline at end of file