Paraleucilla solangeae, Cavalcanti, Fernanda F., Menegola, Carla & Lanna, Emilio, 2014

Cavalcanti, Fernanda F., Menegola, Carla & Lanna, Emilio, 2014, Three new species of the genus Paraleucilla Dendy, 1892 (Porifera, Calcarea) from the coast of Bahia State, Northeastern Brazil, Zootaxa 3764 (5), pp. 537-554 : 539-544

Paraleucilla solangeae sp. nov.

Etymology. For Dr. Solange Peixinho, in recognition of her dedication to the taxonomy of calcareous sponges, mainly species from Bahia and other states in northeastern Brazil.

Diagnosis. Paraleucilla with a single apical osculum ornamented by a short fringe formed by trichoxea and small sagittal tetractines. Tufts of diactines are positioned perpendicular to the sponge surface, crossing two thirds of the choanosome. They are regularly distributed (i.e., equally spaced) along the sponge skeleton. The cortical skeleton is composed of giant tetractines. Both the outer and inner regions have a single type of sagittal triactines and tetractines. The atrium is lined with a tangential layer of sagittal triactines and tetractines.

Type material. Holotype: UFBA POR 3525 [Guarajuba (12°44’12.6’’S 38°05’12.4’’W), Camaçari, Bahia, Brazil; depth: 23 meters; --/ II/2008], Paratypes: UFBA POR 4242 [Guarajuba (12°44’12.6’’S 38°05’12.4’’W), Camaçari, Bahia, Brazil; depth: 23 meters; --/ II/2008], and UFBA POR 4243 [Guarajuba (12°44’12.6’’S 38°05’12.4’’W), Camaçari, Bahia, Brazil; depth: 23 meters; --/ II/2008].

Type locality. Guarajuba, Camaçari, Bahia, Brazil.

Description. The body shape varies from spherical to tubular. The holotype is the largest specimen, and it measures 8.0 x 8.0 mm (osculum-basis axis and width, respectively) ( Figure 2 A). The color when alive is unknown, but the ethanol-preserved specimens are beige. The surface is smooth, but this is most likely because the tips of the diactines are broken (see description below). There is a single apical osculum, which in the holotype (UFBA POR 3525) and in one of the paratypes (UFBA POR 4242) is ornamented by a fringe of trichoxea ( Figure 2 B). Small sagittal tetractines are present at the base of this fringe, and they are covered by the giant spicules that form the cortical skeleton. The body wall is thick in the holotype, which has the narrowest atrial cavity. The paratypes present a wide atrial cavity. The aquiferous system is leuconoid, with the rounded choanocyte chambers typical of the genus.

There are diactines perpendicular to the sponge surface, crossing two thirds of the choanosome (from the outer region to the inner region). They are organized in tufts that are regularly arranged in the sponge body ( Figures 2 C, D). The cortical skeleton is composed of the basal system of giant tetractines ( Figures 2 C, E), which point their apical actine toward the atrium. The inarticulated skeleton (i.e., the outer region) is formed by these apical actines and by the unpaired actines of subatrial triactines and tetractines. Both categories are common in the subatrial skeleton, and they also form the inner region, where the spicules are irregularly distributed. The atrial skeleton is composed of tangential triactines and tetractines. The latter project their apical actines into the atrial cavity ( Figures 2 F, G).

Spicules. ( Table 1). Cortical tetractines ( Figure 3 A): Giant spicules. All actines are conical and have sharp tips. In the basal system, the unpaired actine is shorter than the paired actines, which are curved. The apical actine is straight. [Paired actines: 200 – 373.6 ± 27.8 – 570/ 35.4 ± 2.6 µm; unpaired actine: 190 – 311.2 ± 61.9 – 520.0/ 30.6 ± 6.2 µm; apical actine: 220 – 500.4 ± 48.2 – 750/ 36.8 ± 1.9 µm (n= 3 specimens)]. Subatrial tetractines ( Figure 3 B): Sagittal. Actines are conical and have sharp tips. The paired actines are straight, whereas the unpaired actine is slightly undulated. In general, the latter is longer than the paired actines. The apical actine is short, smooth, and curved. [Paired actines: 160 – 359.6 ± 8.1 – 560/ 35.3 ±4.0 µm; unpaired actine: 170 – 403.3 ± 17.2 – 650/ 36.8 ± 4.2 µm; apical actine: 70 – 182.3 ±98.4 – 430.0/ 26.8 ± 6.1 µm (n= 3 specimens)]. Subatrial triactines ( Figure 3 C): Sagittal. Actines are conical and have sharp tips. In general, the unpaired actine is longer than the paired actines, which are curved. In one of the paratypes (UFBA POR 4242), these spicules have a wide range of sizes. We confirmed that they comprise a single category by plotting a histogram ( Figure 4 A). [Paired actines: 110 – 268.4 ± 57.0 – 550/ 16.5 ± 5.9 µm; unpaired actine: 170 – 412.4 ± 59.2 – 630/ 18.9 ± 5.8 µm (n= 3 specimens)]. Atrial tetractines ( Figure 3 D): Thinner than any other spicule category. Actines are cylindrical and straight (with the exception of the apical one), and they have sharp tips. The apical actine is short and curved close to its tip. [Paired actines: 110 – 223.0 ± 30.7 – 350/ 9.0 ± 2.1 µm; unpaired actine: 100 – 260.0 ± 24.0 – 510/ 10.4 ± 0.4 µm; apical actine: 50 – 104.1 ± 14.6 – 170/ 9.1 ± 1.2 µm (n= 3 specimens)]. Atrial triactines ( Figure 3 E): Sagittal, actines are conical and have sharp tips. The paired actines are curved and are longer than the unpaired one, which is straight. [Paired actines: 90 – 148.8 ± 7.4 – 280/10.5 ± 1.2 µm; unpaired actine: 40 – 88.6 ± 5.2 – 170/ 12.9 ± 0.6 µm (n= 3 specimens)]. Diactines ( Figures 3 F, G): Straight and fusiform. Both tips are sharp, but whereas the proximal one (located inside the sponge) thins gradually ( Figure 3 F), the distal tip (located outside the sponge) comes abruptly to a point ( Figure 3 G). Frequently, this spicule has its distal tip broken. [160 – 498.0 ± 103.7 – 850/ 14.3 ± 2.8 µm (n= 3 specimens)].

Ecology. All the analyzed specimens were sampled by dredging during the monitoring of a submarine emission pipe on the North coast of Bahia (see Materials and Methods above). Specimen UFBA POR 3525 (the holotype) is attached to a fragment of an organism that is most likely a soft coral, and hydroids, calcareous algae, sand, and foraminiferans are also found on its basal surface. On the other specimens (the paratypes), only calcareous algae are observed.

Spicule/ Specimens Actine Length (µm) Width (µm) N

Mean SD Mean SD Cortical tetractines

UFBA POR 3525 (H) paired 381.3 75.3 38.3 9.9 30 UFBA POR 4242 (P) 396.7 82.3 33.5 8.9 30 UFBA POR 4243 (P) 342.7 96.0 34.3 9.4 30 UFBA POR 3525 (H) unpaired 267.5 36.0 26.2 4.8 0 4 UFBA POR 4242 (P) -- -- -- -- -- UFBA POR 4243 (P) 355.0 233.3 35.0 21.2 0 2 UFBA POR 3525 (H) apical 512.7 114.2 38.7 8.5 30 UFBA POR 4242 (P) 541.3 146.3 34.8 7.9 30 UFBA POR 4243 (P) 447.3 92.7 36.9 8.4 26 Subatrial tetractines

UFBA POR 3525 (H) paired 365.3 83.9 37.3 9.7 30 UFBA POR 4242 (P) 350.3 100.3 30.7 9.1 30 UFBA POR 4243 (P) 363.3 95.6 38.0 10.3 15 UFBA POR 3525 (H) unpaired 412.7 91.9 37.5 8.3 30 UFBA POR 4242 (P) 413.7 120.6 32.2 9.3 30 UFBA POR 4243 (P) 383.5 98.8 40.6 11.6 17 UFBA POR 3525 (H) apical 223.3 68.1 31.7 7.6 0 3 UFBA POR 4242 (P) 253.7 96.2 28.7 6.4 0 8 UFBA POR 4243 (P) 70.0 -- 20.0 -- 0 1 Subatrial triactines

UFBA POR 3525 (H) paired 248.7 67.1 15.5 4.8 30 UFBA POR 4242 (P) 332.7 77.1 22.8 6.8 30 UFBA POR 4243 (P) 224.0 84.4 11.2 3.6 20 UFBA POR 3525 (H) unpaired 382.7 97.2 17.7 6.0 30 UFBA POR 4242 (P) 480.7 81.7 25.2 6.6 30 UFBA POR 4243 (P) 374.0 77.1 13.7 3.7 30 Atrial tetractines

UFBA POR 3525 (H) paired 239.7 46.8 9.0 2.3 30 UFBA POR 4242 (P) 241.7 46.8 11.2 4.9 30 UFBA POR 4243 (P) 187.5 39.2 6.9 2.6 0 8 UFBA POR 3525 (H) unpaired 277.0 97.0 10.1 2.5 23 UFBA POR 4242 (P) 243.0 94.2 10.7 3.4 30 UFBA POR 4243 (P) -- -- -- -- -- UFBA POR 3525 (H) apical 87.9 24.4 7.8 2.6 29 UFBA POR 4242 (P) 116.3 21.6 10.3 3.2 30 UFBA POR 4243 (P) 108.2 34.0 9.1 7.3 11 Atrial triactine

UFBA POR 3525 (H) paired 144.0 30.2 11.0 2.6 20 UFBA POR 4242 (P) 157.3 36.0 11.3 3.7 30 ......continued on the next page Spicule/ Specimens Actine Length (µm) Width (µm) N

Mean SD Mean SD

UFBA POR 4243 (P) 145.0 30.4 9.1 7.3 11 UFBA POR 3525 (H) unpaired 90.0 32.4 13.0 3.4 20 UFBA POR 4242 (P) 93.0 34.8 13.5 3.5 30 UFBA POR 4243 (P) 82.8 23.0 12.3 7.8 30 Diactines*

UFBA POR 3525 (H) 499.6 129.3 15.2 3.6 30 UFBA POR 4242 (P) 601.0 111.5 16.5 3.9 30 UFBA POR 4243 (P) 393.5 87.8 11.1 4.5 14 Remarks. Paraleucilla solangeae sp. nov. is morphologically similar to other species of the genus Paraleucilla occurring in the SW Atlantic along the South American coast. It shares with P. perlucida Azevedo & Klautau, 2007 the presence of tufts of diactines. The tufts of diactines in P. perlucida are also regularly spaced ( Klautau 2013, personal communication), as we observed in P. s o l a n g e a e sp. nov. Nevertheless, these species can be easily distinguished, mainly by the sizes of their spicule categories, by the layer of tangential triactines that is present only in P. perlucida , and by the composition of the atrial skeleton, which is formed by triactines and tetractines in P. solangeae sp. nov. but is formed exclusively by tetractines in P. perlucida . Moreover, in P. perlucida , a second subatrial skeleton is occasionally present.

Paraleucilla sphaerica Lanna et al., 2009 also resembles P. solangeae sp. nov. Here are the main differences between these species: the presence of microdiactines [94.5 – 177.5 ± 5.8 – 315/ 9.0 ± 1.2 µm] in the cortical and choanosomal skeleton of P. sphaerica , the presence of diactines [160 – 498.0 ± 103.7 – 850/ 14.3 ± 2.8 µm] arranged in tufts in P. solangeae sp. nov., and the abundance of subatrial tetractines in P. solangeae sp. nov., which are comparatively rare in P. sphaerica .

The tufts of diactines also represent a key difference between P. solangeae sp. nov. and the two new species described here (see below).

Figures

FIGURE 2. Paraleucilla solangeae sp. nov. A—Preserved holotype (UFBA POR 3525); B—Detail of the oscular fringe; C— Transverse section [from the cortex to the atrium] showing the outer region (or) and inner region (ir). Arrows indicate the presence of tufts of diactines; D—Detailed view of a tuft of diactines (arrow) inserted close to the cortex (cx); E—Tangential section showing the cortical skeleton; F—Tangential section showing the atrial skeleton and the apical actines (arrows) of some atrial tetractines; G—Detailed view of the apical actines of tetractines pointed toward the atrial cavity. (B, C, E, G: UFBA POR 3525, holotype; D, F: UFBA POR 4242, paratype).

FIGURE 3. Spicules of Paraleucilla solangeae sp. nov. A—Cortical tetractine; B—Subatrial tetractine; C—Subatrial triactine; D—Atrial tetractine; E—Atrial triactine; F—Diactine (distal tip is broken); G—Detailed view of the distal tip of a diactine. P-paired actines; u- unpaired actine.

FIGURE 4. Histograms and box plots of the spicule measurements for the three new species: A— P. solangeae sp. nov.; B – E— P. o c a sp. nov. and F-H: P. incomposita sp. nov. A—Histograms showing the distributions of actine length for the subatrial triactines of the paratype of P. solangeae sp. nov. (UFBA POR 4242; top panel: paired actines, bottom panel: unpaired actines). B – C—Boxplots of the categories of subatrial triactines of P. oca sp. nov. (B—paired actines and C—unpaired actines). D – E— Boxplots of the categories of subatrial tetractines of P. o c a sp. nov. (D—paired actines and E—unpaired actines). F – H— Boxplots of the categories of subatrial triactines of P. incomposita sp. nov. (F—paired actines; G—unpaired actines and Hapical actines). Each box displays the median and upper and lower quartiles of the spicules measurements for each category. Box whiskers represent the maximum and minimum range. The p-value indicates a significant difference between spicule categories.