The genus Clathria from the Gulf of Mexico and Mexican Caribbean, with redescription and resurrection of Clathria carteri (Poecilosclerida: Microcionidae) Author Gómez, Patricia text Zootaxa 2014 3790 1 51 85 journal article 36866 10.11646/zootaxa.3790.1.3 6e418488-0a69-434a-872d-bbd01345d08d 1175-5326 285913 CB58F85A-924D-4148-AAC4-CDBD041EB3CD Clathria ( Clathria ) foliacea Topsent, 1889 ( Figs. 2 –4, 7–8, 20A, Tab. 1 ). Clathria foliacea Topsent, 1889 :39 . Thalyseurypon foliacea sensu de Laubenfels 1936a :107 . Clathria ( Clathria ) foliacea sensu Hooper 1996 :173 . Non Clathria carteri Topsent, 1889 :38 (= Clathria ( Clathria ) carteri ). Species status : The holotypes designated by Topsent (1889) are missing, including those of other Clathria species, viz. C. foliacea , C. carteri , C. fascicularis , C. copiosa and C. dentata from the Campeche Bank (in van Soest 1984 and I. Domart-Coulon pers. comm.). The only comparative material available is a fragment ( 10 mm 2) of Thalyseurypon foliacea (NHMN 22516) from de Laubenfels (1936a) kindly sent on loan by Drs. K. Rützler and W. Moser. Another attempt to trace type material of microcionids was by Dr. Hooper, who was unable to find several of the types for his monograph on microcionids (1996) (pers. comm.). Therefore, a neotype is designated here from the type locality (Campeche Bank, Mexico ), and a comprehensive redescription of the species is provided to clarify the features and validity of this species. Neotype (here designated): CNPGG–1178 Campeche Bank, 2/VIII/1996 , 78 m depth, bushy habit, red when alive. FIGURE 2. Different morphs of Clathria ( Clathria ) foliacea Topsent, 1889 : A. CNPGG–756 Campeche Bank. B. USNM 22516 Tortugas, Florida. C. CNPGG–064 Yucatan Channel. D. CNPGG–1188 Quintana Roo. E. CNPGG–1201 Juvenile specimen, Quintana Roo. F. CNPGG–1195 Campeche. G. NEOTYPE CNPGG–1178 Campeche Bank. Scale bars: A, 5 cm; B, C, F, G, 2.5 cm; D, E, 1 cm. FIGURE 3. Clathria ( Clathria ) foliacea Topsent, 1889 : A. Skeletal pattern from a flabellate branch. B. Detail of a plumose tract. Scale bars: A, B, 200 µm. FIGURE 4. SEM depiction of spicules from Clathria ( Clathria ) foliacea Topsent, 1889 : A-A1. (Sub) tylostyles and detail of tuberculated head. B-B1. Thin styles and microspined head. C. Acanthostyles, scantily spined shaft, coarse tuberculated head. D. Toxa in different shapes. E. Twisted palmate isochelae. Scale bars: A, 100 µm; B, D, 50 µm; A1 B1, 5 µm; C, 40 µm; E, 2.5 µm. Description of neotype . A bushy sponge arising from a short, thin stem. The whole sponge is a cluster of foliaceous branches anastomosed and bifurcated with jagged edge, up to 10 cm in total height, 7.5 cm in upper diameter, the leaves are 2 mm thick, the stem is 15 mm thick, 25 mm long ( Fig. 2 G). It is dull red when alive. Under the microscope it has a micro-hispid surface and neither oscules nor pores were seen. Choanosomal skeleton is characterized by a plumo-reticulation axially displayed of thick styles echinated by acanthostyles; the ectosome is a thin sheet composed of slender auxiliary styles disposed in a perpendicular plane at the surface and some others tangentially disposed with twisted palmate isochelae, and toxas (Spicule measurements Tab. 1 ). Material examined. USNM 1156280 Campeche Bank 19º45’12”N 91º46’18”W , 6/IV/1983 , 47 m depth, bushy habit. CNPGG –064 Yucatan Channel 21º58’N 86º43’W , 26/X/1985 , 40 m depth, flabellate branches. CNPGG –351 Madagascar reef, Yucatan 21º26'16.6"N 90º16'39"W , 13 m depth, red labyrinth shape. CNPGG –489 Yucatan Channel 22º2’4”N 87º2’5”W , 25/IV/1985 , 35 m depth, bushy habit. CNPGG –756 Campeche Bank 19º45’12”N 91º46’18”W , 6/IV/1983 , 47 m depth, carbonate–mud bottom, flabellate bush habit. CNPGG –1185 Campeche Bank 19º45’12”N 91º46’18”W , 6/IV/1983 , 47 m depth, flabellate bush habit. CNPGG –1187 Campeche Bank 22°30’6’’N 89°00’27.18’’W , 15/XI/2005 , 25 m depth bushy habit pinkish red alive. CNPGG –1188 Puerto Morelos, Quintana Roo 20º50’36.91”N 86º52’ 27.85W , 25/IV/1979 , 12 m depth, red labyrinth shape. CNPGG –1189 Campeche Bank 19º45’12’’N 91º46’18”W , 6/IV/1983 , 47 m depth, carbonate–mud bottom, bushy habit. CNPGG –1195 Campeche, Campeche 19º58.387”N 90º59.752”W , 7/IX/2003 , 59 m depth, flabellate bush habit. CNPGG –1196 Madagascar reef Yucatan, 21º26’28.3”N 90º17’34.0”W , 11/VII/2011 , 6 m depth. CNPGG –1201 Xahuayxol, Quintana Roo, 24/IV/1992 , flabellate habit. CNPGG –1224 Progreso Yucatan 21º16’49.55”N 89º42’23.16”W , 8/X/2009 , 1 m depth. CNPGG –1228 Serpientes reef, Yucatan 21º26’24”N 90º28’25.39”W 13/VI/2011 , 15 m depth, bushy habit. CNPGG –1242 Isla Sacrificios Veracruz 19º10’30”N 95º6’W , XII/1981 , flabellate bush habit. CNPGG –1358 Madagascar reef, Yucatan 21º26'16.6"N 90º16'39"W , 7/VI/ 2011 , 10 m depth, scarlet, labyrinth shape. CNPGG –1404 Campeche Bank 21º6.60’N 92º 8.66W , 9/VI/2005 , 54 m depth. CNPGG –1416 Bacachat, Yucatan 21º26’20.90”N 90º16’58.90”W , 4/VI/2012 , 12 m depth, rocky substrate. Comparative material examined. A fragment from Thalyseurypon foliacea sensu de Laubenfels (1936) ( USNM 22516), south of Loggerhead Key, Tortugas , Fla. 28/VII/ 1932 , 105 m depth. Samples of Clathria ( Clathria ) carteri from the present study (see below). Description. The present species is bushy, generally with flabellate branches. Sometimes the branches are anastomosed, labyrinthine, sometimes with independent limbs; they arise from a narrow and short stem which supports a rosette of leaves outlining a shrub. In one case, branches grow without a stem from an encrustation over a bivalve conch (CNPGG–1188). The total height is 5–24 cm , upper diameter 3–27 cm , stem length 6–40 mm , stem diameter 10–40 mm , and thickness of the flabellate branches 1–3.2 mm ( Fig. 2 A–G). A juvenile specimen is represented by a single frond 10 mm at the base, 45 mm high ( Fig. 2 E). The most prevalent color when alive is bright red or dull red, but pinkish cherry can be found; in spirit and when dry it is beige or drab. The consistency is resilient, slightly compressible. The surface is microscopically hispid owing to the projection of plumose styles. Oscules or pores are not visible. The dermal layer consists of auxiliary styles, palmate isochelae, and toxas sustained by the protruded styles delineating the surface. Skeleton. ( Fig. 3 A–B) The choanosomal skeleton is an irregular reticulation but clearly distinguished by strongly developed ascending fibers characterized by an amber color, with profusely plumose thick styles along the fiber with points directed outward, and sparsely echinated by acanthostyles. These fibers are usually cored by thick styles, some acanthostyles may be included also, in places they are free. Fiber diameter ranges from 30–70 µm, few of them 100–350 µm, but this is difficult to measure since the plumose styles among echinators and other single spicules dispersed all around the flesh obscure the spongin line of the fiber. Connective fibers are free from spicules, short, and of 10–50 µm in diameter. Samples 0 64 and 1187 have no clear distinction between fibers, since they measure the same in diameter. The ectosome consists of echinated thick styles as well as those in plumose tracts projecting outside the surface, which are supported by ascendant tracts, in addition, single acanthostyles are echinating on fibers parallel to the surface. Spicules. (Fig. 4A–E) Auxiliary styles to subtylostyles slender and straight shaft, sometimes sinuous with smooth and microspined heads: 101.4–441.5 ×1.3–8.3 µm. Tylostyles to subtylostyles thick, straight, slightly bent with tuberculated or bumpy heads: 122.2–540 × 7.8–38 µm. Acanthostyles with sparse spines along the shaft, sometimes minutely spined at the end of the shaft, sometimes smooth, but coarsely tuberculated head: 66.5–124.8 × 3.9–9.8 µm. Palmate isochelae always twisted, 7.2–11.7 µm. Wing-shaped toxas recurved and with shallow arms 13–140.4 µm. The two microscleres are occasionally rare and sometimes absent (Measurements in Tab. 1 ). Remarks. Clathria ( Clathria ) foliacea shows some intraspecific morphological variability, which at first sight was thought to indicate different species, but thorough examination (skeleton and spiculation) showed specimens to be conspecific. The present samples of C. ( C. ) foliacea nearly all have the consistent presence of microscleres (isochelae and toxas), in addition to features that characterize the species. These microscleres are not mentioned in Topsent’s work; probably because the presence of microscleres, specially of isochelae, are extremely rare in some specimens or difficult to find, as was the case of Thalyseurypon foliacea and specimen CNPGG–1195 (the latter with only two isochelae and very few toxas in the field) studied herein. However, they proved to be quite similar in the skeletal arrangement, as well as in geometry and the spicular measurements to the other samples compared ( Tab. 1 ). Accordingly, samples were allocated to C. ( C. ) foliacea . TABLE 1 . Spicule micrometries, distribution/depth (in meters) of Clathria ( Clathria ) foliacea Topsent 1889 . Measurements are in µm, mean values are in bold, measurement number n=30. YUC, Yucatan Channel; C.B., Campeche Bank; Q.R., Quintana Roo; Cam, Campeche; Fla, Florida; –, Absent; n.r., not recorded. (* Thalyseurypon foliacea de Laubenfels’s material re-measured); (♠ Neotype designated).

Specimen	Location	Style I	Style II	Acanthostyle	Toxa Isochela
CNPGG	YUC/	109.2– 246.2 –430/	160.8– 318.4 –540/	80.6– 99.7 –115.1/	33.8– 90.56 –140 7.2– 9.6 –12
64	40	2– 4.46 –7.8	10.9– 20.25– 32.2	5.2– 6.4 –9.8
CNPGG	YUC/	113– 246.8 –430/	156– 315.5 –423/	87.8 –104.2 –119/	52– 85.9 –135.2 7.8– 8.9 –10
489	35	2– 4.7 –7.8	10– 16.8 –25.7	5.2– 7.6 –9.1
CNPGG	C.B./	101.4– 241.3 –410/	166– 322.6 –477/	66.5– 96.6 –119.6/	26– 83 –125.3 8.5– 9.7 –11
756	47	2– 4.54 –8.3	9.1– 20.1– 31.2	5.2– 6.68 –8.3
CNPGG	C.B./	121.6– 229.2 –350/	158– 290.5 –420/	72.5– 100.4 –124.8/	31.3– 53.7 –83.2 7.3– 9 –10.4
1185	47	2– 4.6 –7.8	13– 23 –36.4	6.7– 7.78 –9.8
CNPGG	C.B./	122.2– 244.7 –415/	184.6– 305.9 –420/	80.6– 96.5 –114.4/	41.6– 60.7 –101 8.3– 9.7 –11
1187	25	2– 4.88 –7.8	9.1– 16.1 –33.8	3.9– 5.8 –7.2
CNPGG	Q.R./	129.7– 266.2 –426/	122.2– 263.9 –435/	72.8– 88 –106.6/	44.2– 71.1 –130 7.8– 10– 11.7
1188	12	2.6– 4.5 –7.8	10– 15 –23.4	5.2– 7.1 –9.6
CNPGG	Cam/	112.3– 256.2 –402/	150– 305.9 –480/	68.9– 89.9 –112/	13–19.8–75.4 8.5–11.7
1195	59	1.3– 4.1–7.8	7.8– 14.8 –21	4.4– 6.13– 7.8
CNPGG	Q.R./	140.6– 256.8 –356/	146– 307.7 –467/	72.6– 95.7 –121.6/	15.1–59.2 7.8– 9.5 –10
1201	n.r.	1.5– 3.9 –7.5	10.1– 16.1 –24.7	5.2– 6.8 –9.3
CNPGG	C.B./	106.6– 227– 441.5/	177.6– 320 –470/	92.5– 99.6 –111/	41.6– 59.1 –138 7.3– 8.3 –12
1178♠	78	2– 4.5 –8	9.1– 24.2 –38	5– 7.4 –8.5
USNM 22516*	Fla/ 105	106.6– 261.5 –437/ 1.5– 3.8 –8	146– 295.8 –435/ 9.1– 15.4 –23.4	70– 94– 119/ 4.9– 5.5 –6.7	– –
Average	246/4.4	304.2/18	96.3/6.7	67.8 9.4

The growth forms of Clathria species elsewhere in the Caribbean, with the exception of that of Florida ( T. foliacea ), do not include the characteristic bladed forms as described by Topsent and in the present work. This strengthens the conspecificity with the present species. A species closely related to C . ( C .) foliacea is C . carteri Topsent (1889) and has even been considered a synonym of C . ( C .) foliacea (de Laubenfels 1936a ; van Soest 1984 ; Hooper 1996 ) because of their close morphology according to the literature, and nobody has had the opportunity to collect it or study it up to now. Five other samples of Clathria sp. were initially assigned to C. ( C .) foliacea , but detailed comparison with the present material indicated they belonged to another species different from C. ( C .) foliacea , and consequently reallocated to C. ( C. ) carteri (see below). Despite their similarity in morphology, the distinctive skeletal pattern as well as the different geometry and size of spiculation leading to the resurrection of C. ( C. ) carteri , these are not conspecific with C. ( C .) foliacea . Distribution. As far as is known, the species has only been found within the Gulf of Mexico : the type locality Campeche and at Loggerhead Key, Fla. Its distribution has been extended from the present study to include the Caribbean at the Yucatan Channel and Quintana Roo, Mexico , at 1–105 m depth.