Tanacetipathes tanacetum ( Pourtalès, 1880 ), Pourtales, 1880

Loiola, Livia L. & Castro, Clovis B., 2005, Tanacetipathes Opresko, 2001 (Cnidaria: Antipatharia: Myriopathidae) from Brazil, including two new species, Zootaxa 1081, pp. 1-31 : 12-19

Tanacetipathes tanacetum ( Pourtalès, 1880)

Figures 7–10

Antipathes tanacetum Pourtalès, 1880: 116 , pl. 3, Fig. 13 ; Brook, 1889: 162; Opresko, 1972: 984 – 986, tab. 2, Fig. 7 ; Echeverría & Castro, 1995: 1 –7, Figs. 2–5 (part).

Tanacetipathes tanacetum: Opresko, 2001a: 358 –361, Figs 11 a and 12d; 2001b: 349, Figures 7–10 ; Pérez et al., 2005: 3 –8, Figs 1–4 .

Material examined. Brazil: off Aracruz: 19 ° 37’ S, 0 38 ° 41’ W, about 65 m, Bahia­1 ( MNRJ 3410: 3 colonies); 19 ° 42’ S, 0 39 ° 26’ W, 239 m, REVIZEE Bahia­2 Sta. #E0533, ( MNRJ 4604, MNRJ 4612, MNRJ 4617, MNRJ 5152: 14 colonies); 19 ° 45’ S, 0 39 ° 31’ W, 60 m, REVIZEE Central V Sta. #27 ( MNRJ 4861, MNRJ 4862: 2 colonies); 19 ° 46’ S, 0 39 ° 29’ W, 498 m, REVIZEE Central VI Sta. # 3 ( MNRJ 4935: 1 colony); off Vitória, 20 ° 27’ S, 0 39 ° 44’ W, 1293 m, REVIZEE Bahia­2 Sta. #E0536 ( MNRJ 4634: 1 colony); Jaseur Bank, 20 ° 36’ S, 0 35 ° 51’ W, about 100 m, REVIZEE Central II ( MNRJ 3697: 6 colonies); off Guarapari: 20 ° 40’ S, 0 34 ° 35’ W, 100 m, REVIZEE Central II Sta. #46 ( MNRJ 3696, MNRJ 5149: 8 colonies); 20 ° 40’ S, 0 34 ° 35’ W, 108 m, REVIZEE Central V Sta. #45 ( MNRJ 4668, MNRJ 4672, MNRJ 5153, MNRJ 5159, MNRJ 5161: 16 colonies); 20 ° 40’ S, 0 35 ° 28’ W, about 60 m, REVIZEE Central II ( MNRJ 3693: 1 colony); Davis Bank, 20 ° 41’ S, 0 37 ° 07’ W, about 57 m, REVIZEE Central IV ( MNRJ 3393: 1 colony); off Guarapari, 20 ° 42’ S, 0 35 ° 27’ W, 60 m, REVIZEE Central V Sta. #21 ( MNRJ 4863: 2 colonies); off Itapemirim, 21 ° 09’ S, 0 40 ° 16’ W, about 100 m, REVIZEE Central Sta. #14 ( MNRJ 2975: 1 colony); off São João da Barra: 21º20’ S, 040º16’ W, 120 m ( MNRJ 2369: 2 colonies); 21º20’ S, 040º15’ W, 183 m ( MNRJ 2363: 1 colony); off Cape of São Tomé: 22 ° 00’ S, 0 40 ° 05’ W, 100 m, REVIZEE Central V Sta. #38 ( MNRJ 4665, MNRJ 5140, MNRJ 5151, MNRJ 5157: 12 colonies); 22 ° 03’ S, 0 40 ° 06’ W, 91 m, REVIZEE Central Sta. #02 ( MNRJ 3018; 3023: 2 colonies); Almirante Saldanha Bank, 22 ° 22’ S, 0 37 ° 35’ W, 103 m, REVIZEE Central VI ( MNRJ 4917, MNRJ 4918, MNRJ 5162: 6 colonies); Almirante Saldanha Bank, 22 ° 22’ S, 0 37 ° 35’ W, 240 m, REVIZEE Central VI ( MNRJ 4927, MNRJ 4928, MNRJ 5160, MNRJ 5163, MNRJ 5164: 24 colonies); off Cabo Frio: 22 ° 44’ S, 0 31 ° 49’ W, 80 m, REVIZEE Central V Sta. #42 ( MNRJ 4670; 4671: 2 colonies); 22º54’ S, 040º48’ W, 156 m ( MNRJ 2353: 1 colony).

Diagnosis. Colonies ( Figs. 7 a–b; 8a; 9a–b; 10a) monopodial (up to 65 cm tall) or rarely with branches up to the 2nd order, emerging near the colony base ( Figs. 7 a–b; 8a; 9a–b; 10a), pinnulated; pinnules arranged biserially in 4–6 longitudinal rows (depending on the presence of second anteriors) and in alternate groups along the axis. Maximum length of anterior primary pinnules 3–19 mm (average 10.03 ± 3.48 mm), maximum length of posterior primaries 6–27 mm (average 14.01± 4.35 mm); posterior primaries curved distally towards the anteriors ( Figs. 7 c, 8b, 9c, 10b). Secondary pinnules up to 22 mm long (average maximum length 12.27± 3.56 mm); 0–4 secondaries on each anterior primary, 1–7 on each posterior primary, usually in single series on the abpolypar side of the primaries, occasionally on the polypar side ( Figs. 7 c, 8c, 9b, 10b). Tertiary pinnules on the abpolypar (rarely polypar—see fig. 9c) side of secondaries and sometimes bearing quaternaries ( Figs. 7 c, 8c, 9b, 10b). Spines ( Figs. 7 d–h; 8d–g; 9c–g; 10c–f) compressed, with acute apex, length 3–4 times the width; polypar spines 0.04–0.27 mm tall, abpolypar 0.01–0.16 mm ( Figs. 7 d–h; 8d–g; 9c–g; 10c–f). Polyps 0.6–0.8 mm in transverse diameter, arranged in a single series, restricted to the polypar side (concave) of pinnules and subpinnules; 10 – 13 per centimeter (emended from Opresko, 1972).

Remarks. The Brazilian specimens were treated as four different morphotypes of Tanacetipathes tanacetum ( Tab. 1 ). These forms were considered as a single species since there are previous records of great variability in T. tanacetum and because we could not find continue gaps in their variation. In the first form ( Fig. 7 ), the length of the primary pinnules and the arrangement of secondaries are as usually observed in T. tanacetum , according to the diagnosis given by Opresko (1972). However, Opresko (1972) indicates two groups of specimens based on the height of polypar spines: (1) up to 0.13 mm tall and (2) more than 0.23 mm tall. The polypar spines’ height in our “first form” varies continuously along the range of 0.12–0.27 mm, as described by Echeverría & Castro (1995). The distribution of polypar spines height varied in the 20 specimens assigned to this form as follows: seven specimens have polypar spines varying along the whole range cited above, 10 have all polypar spines smaller than 0.20 mm, and three have all polypar spines taller than 0.20 mm.

The colonies of the second form ( Fig. 8 ) are similar to the group of specimens with smaller spines referred to by Opresko (1972: 0.10–0.13 mm tall). Besides, these colonies have primary pinnules with lengths a little smaller (18 mm) than that described by Opresko (1972—up to 25.0 mm) and by Echeverría & Castro (1995—up to 20.5 mm). Tanacetipathes spinescens ( Brook, 1889) shows similar values for these characters (primary pinnules up to 15.0 mm; spines up to 0.19 mm), but this species has densely branched colonies ( Brook, 1889; Opresko, 2001a). All forms of T. tanacetum are sparsely branched, only up to the second order, and usually branches develop from near the base.

The third form ( Fig. 9 ) differs from the first two especially in the presence of fewer and smaller secondary pinnules ( Tab. 1 ), but relatively large spines (up to 0.27 mm). Opresko (1972) reported reduced subpinnulation in some colonies of T. tanacetum , in a condition comparable to that observed in T. hirta . However, this form differs from T. hirta because the latter has small spines (up to 0.13 mm tall) and branching up to the 5th order.

The subpinnulation in the fourth form ( Fig. 10 ) is also reduced, as in the third form and in T. hirta . The spines’ heights are also similar to T. hirta spines in the descriptions of Opresko (1972­polypar 0.07–0.13 mm [see diagnosis], abpolypar 0.03–0.10 mm), and Warner (1981­polypar 0.09–0.17 mm, abpolypars 0.04–0.13 mm), and in the description of this species herein included. However, specimens of this fourth form are branched only up to the 1st order, while T. hirta is branched up to the 5th order.

A lectotype for this species was chosen and described by Pérez et al. (2005), because the type­series of T. tanacetum , designated by Pourtalès (1880), is heterogeneous. According to these authors, based on the length of the primary pinnules, and the number and arrangement of the secondary pinnules, there appear to be as many as four different species represented in the type­series. The spines size was not considered a sound character to differentiate these four morphotypes mentioned, as this character varies substantially from specimen to specimen ( Pérez et al., 2005).

The specimen chosen as lectotype has posterior primary pinnules up to 1.2 cm long. However, according to Pérez et al. (2005), they are usually up to 2.0 cm and rarely up to 2.5 cm. Among the 89 Brazilian specimens examined herein, 36 have posterior primaries at most 1.2 cm long, 44 have the longest posterior primaries between 1.2 and 2.0 cm, and 9 specimens have posterior primaries that exceed 2.0 cm long. The overall maximum length of posterior primaries reaches 2.7 cm.

In the lectotype description, there are one to three secondary pinnules on the posterior primary pinnules, usually arranged uniserially on the abpolypar side. However, the illustration given by Pérez et al. (2005: Fig. 2 b) shows four to five secondaries per posterior primary, arranged both in the polypar and abpolypar sides of the pinnules. Of the 89 Brazilian specimens considered in this study, 42 have up to three secondaries per posterior primary pinnules, 40 have four or five, and seven have more than five. The maximum observed is up to seven. Considering these characteristics, Brazilian specimens are very similar to the lectotype of T. tanacetum designated by Pérez et al. (2005).

Opresko’s (1972) diagnosis was emended to accommodate variations of the four different forms of the Brazilian specimens mentioned above.

Distribution. Atlantic: “Lesser Antilles” ( Pourtalès, 1880; Brook, 1889; Opresko, 1972). Brazil: off the Parcel do Manoel Luís (00°17’ N) ( Opresko, 1972); off the Atol das Rocas (about 03°50’ S) ( Opresko, 1972); off eastern Brazil (19°37’–23° S) ( Echeverrìa & Castro, 1995; current records—Fig. 1).

Figures

FIGURE 1 — Distribution of Tanacetipathes Opresko, 2001 off Brazil, between 13 ° – 22 ° S: T. barbadensis (Brook, 1889); T. tanacetum (Pourtalès, 1880); T. hirta (Gray, 1857); T. thamnea (Warner, 1981); T. longipinnula n. sp.; T. thalassoros n. sp.

FIGURE 2 — Schematic draw of a colony, with some of the main terms used on descriptions and diagnosis of Tanacetipathes species.

FIGURE 3 — Transversal view of a branch, with the main terms used in the descriptions and diagnosis of the species of Tanacetipathes: P = polypar side; A = abpolypar side; 1 = first anterior primary pinnule; 2 = second anterior primary pinnule; 3 = posterior primary pinnule; 4 = polypar secondary pinnule; 5 = abpolypar secondary pinnule; 6 = polypar tertiary pinnule; 7 = abpolypar tertiary pinnule; 8 = polypar spines; 9 = abpolypar spines.

FIGURE 7 — Tanacetipathes tanacetum (Pourtalès, 1880), Form 1 (MNRJ 4604): a—Colony; b—Pinnulate region of the colony; c—Transversal view, cycle of pinnules; d—Spines of the basal region of a pinnule; e—Spines of the middle region of a pinnule; f—Spines of the apical region of a pinnule; g—Polypar spines of the middle region of a pinnule, close view; h—Abpolypar spines of the middle region of a pinnule, close view. Scale bars: a = 2.0 cm; b = 1.0 cm; c = 0.5 cm; d­f = 0.2 mm; g = 0.05 mm; h = 0.02 mm.

FIGURE 8 — Tanacetipathes tanacetum (Pourtalès, 1880), Form 2 (MNRJ 4672): a – Colony; b—Transversal view, cycle of pinnules; c—Polypar spines of the middle region of a pinnule, close view; d—Abpolypar spines of the middle region of a pinnule, close view; e—Spines of the basal region of a pinnule; f—Spines of the middle region of a pinnule; g—Spines of the apical region of a pinnule. Scale bars: a = 2.0 cm; b = 0.3 cm; c­d = 0.02 mm; e­g = 0.2 mm.

FIGURE 9 — Tanacetipathes tanacetum (Pourtalès, 1880), Form 3 (MNRJ 4665): a—Colony; b—Pinnulate region of the colony; c—Transversal view, cycle of pinnules; d—Polypar (p) and abpolypar (ab) spines of the middle region of a pinnule, close view; e—Spines of the basal region of a pinnule; f—Spines of the middle region of a pinnule; g—Spines of the apical region of a pinnule. Scale bars: a = 1.0 cm; b = 1.0 cm; c = 0.3 cm; d = 0.05 mm; e – g = 0.2 mm.

FIGURE 10 — Tanacetipathes tanacetum (Pourtalès, 1880), Form 4 (MNRJ 4928): a—Colony; b—Transversal view, cycle of pinnules; c—Polypar and abpolypar spines of the middle region of a pinnule, close view; d—Spines of the basal region of a pinnule; e—Spines of the middle region of a pinnule; f—Spines of the apical region of a pinnule. Scale bars: a = 2.0 cm; b = 0.3 cm; c = 0.05 mm; d­f = 0.2 mm.

FIGURE 11 — Tanacetipathes thamnea (Warner, 1981) (MNRJ 4669): a—Colony; b—Pinnulate region of the colony; c—Transversal view, cycle of pinnules; d—Polypar (p) and abpolypar (ab) spines of the apical region of a pinnule, close view; e—Spines of the basal region of a pinnule; f—Spines of the middle region of a pinnule; g—Spines of the apical region of a pinnule; h—Polypar spines of the middle region of a pinnule, close view; i—Polypar spines of the apical region of a pinnule, close view; j—Abpolypar spines of the middle region of a pinnule, close view. Scale bars: a = 2.0 cm; b = 1.0 cm; c = 0.3 cm; d, h – i = 0.05 mm; e – g = 0.2 mm; j = 0.02.

FIGURE 13 — Maximum number of secondary pinnules per posterior primary pinnule in 49 colonies of Tanacetipathes thamnea from Brazil.

Tables

TABLE 1 — Tanacetipathes tanacetum (Pourtalès, 1880): comparison of morphotypes. hcol = maximum colony height; lap = primary anterior pinnule length; ps / pp = ratio number of secondary pinnules per primary; sppol = height of polypar spines (range); spabpol = height of abpolypar spines (range).

	hcol (cm)	lap (mm)	ps/pp	sppol (mm)	spabpol (mm)
form 1	up to 65	up to 27	up to 7:1	0.12–0.27	0.01–0.16
form 2	up to 45	up to 18	up to 6:1	0.06–0.16	0.01–0.10
form 3	up to 43	up to 20	up to 4:1	0.10–0.27	0.02–0.15
form 4	up to 63	up to 26	up to 4:1	0.04–0.16	0.01–0.10