Cottarellicaris sanctiangeli Bruno & Cottarelli, 2020

Bruno, Maria Cristina, Cottarelli, Vezio, Marrone, Federico, Grasso, Rosario, Stefani, Erika, Vecchioni, Luca & Spena, Maria Teresa, 2020, Morphological and molecular characterization of three new Parastenocarididae (Copepoda: Harpacticoida) from caves in Southern Italy, European Journal of Taxonomy 689, pp. 1-46 : 9-19

Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov.

urn:lsid:zoobank.org:act:BFD63C80-116A-49C7-B10D-547D0146CB5A

Figs 2 – 8 ; Tables 1–3

Diagnosis

Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov. is characterized by the peculiar morphology of the male P4 enp, with apically curved inner tip and, mainly, with a thin proximal outgrowth as long as the apical (crenulate) lamellar outgrowth. Females are characterized by the P3 enp being shorter than the corresponding exp-1.

Etymology

The species epithet is the masculine genitive of the Latin words ‘ sanctus ’ and ‘ angelus ’, meaning ‘Saint’ and ‘angel’, respectively, and refer to the locus typicus of this Cottarellicaris , i.e., a cave dedicated to Saint Michael the Archangel, whose cult is widespread in Southern Italy.

Material examined

Holotype

ITALY • ♂; Cosenza Province, Cassano allo Ionio , Complesso Grotte di Sant’Angelo , Grotta Superiore di Sant’Angelo , pool 2; approximate coordinates 39°47′20.24ʺ N, 16°18′25.60ʺ E; 9 May 2017; R. Grasso and M.T. Spena leg.; dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli holotype: male”; NHMUK 2020.4 .

Paratypes

ITALY • 2 ♂♂; same collection data as for holotype; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: male”; NHMUK 2020.5 to 2020.6 • 2 ♂♂; same collection data as for holotype but pool 4; dissected and mounted each on one slide labelled “ Cottarellicaris sanctiangeli paratype: male”; NHMUK 2020.7 to 2020.8 • 1 ♂; same collection data as for holotype but pool 4; undissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: male”; NHMUK 2020.9 • 3 ♀♀; same collection data as for holotype; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: female”; NHMUK 2020.10 to 2020.12 • 1 ♀; same collection data as for holotype; mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: female”; NHMUK 2020.13 • 1 ♀; same collection data as for holotype but pool 4; dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: female”; NHMUK 2020.14 • 1 ♀; same collection data as for holotype but pool 4; mounted on one slide labelled “ Cottarellicaris sanctiangeli paratype: female”; NHMUK 2020.15 .

Other material

ITALY • 1 ♂; Cosenza Province, Cassano allo Ionio , Vucco Ucciardo , pool 1; approximate coordinates; 39°47′3.56ʺ N, 16°19′14.87″ E; 25 Apr. 2013; R. Grasso and M.T. Spena leg.; mounted on one slide labelled “ Cottarellicaris sanctiangeli : male”; NHMUK 2020.16 • 4 ♂♂; same collection data as for preceding; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli : male”; NHMUK 2020.17 to 2020.20 • 1 ♀; same collection data as for preceding; mounted on one slide labelled “ Cottarellicaris sanctiangeli : female”; NHMUK 2020.21 • 4 ♀♀; same collection data as for preceding; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli : female”; NHMUK 2020.22 to 2020.25 • 2 ♂♂; Cosenza Province, San Lorenzo Bellizzi, Grotta del Banco di ferro , pool 3; approximate coordinates: 39°52′26.38ʺ N, 16°20′27.35ʺ E; 27 May 2015; Grasso and M.T. Spena leg.; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli : male”; NHMUK 2020.26 to 2020.27 • 1 ♂, 1 ♀; same collection data as for preceding; mounted on one slide labelled “ Cottarellicaris sanctiangeli : male and female”; NHMUK 2020.28 to 2020.29 • 2 ♀♀, same collection data as for preceding; each dissected and mounted on one slide labelled “ Cottarellicaris sanctiangeli : female”; NHMUK 2020.30 to 2020.31 • 1 ♀, same collection data as for preceding; 10 May 2017; mounted on slide labelled “ Cottarellicaris sanctiangeli : female”; NHMUK 2020.32 .

Description

Adult male

BODY. Unpigmented, nauplius eye absent. Total body length, measured from tip of rostrum to posterior margin of caudal rami (excluding caudal setae) from 295 to 333 µm, mean 311 μm (n = 6). Habitus ( Fig. 2a ) cylindrical and slender, without any demarcation between prosome and urosome; prosome to urosome ratio: 0.90. Free pedigerous somites without any lateral or dorsal expansions, all connected by well-developed arthrodial membranes.Integument weakly sclerotized, without cuticular pits, ornamented with sensilla on all somites except preanal one. Cuticular windows on urosomites and cephalothorax not observed. Cephalothorax representing about 20 % of total body length. Anal somite ( Fig. 3a, c ) with pair of large dorsal sensilla at base of anal operculum, pair of cuticular lateral pores (one pore on each side) on proximal margin. Anal operculum ( Fig. 3a, c ) well-developed, ornamented with row of spinules on outer surface, with straight distal margin. Anal sinus wide open.

CAUDAL RAMI ( Fig. 3a, c ). Shorter than anal somite, approximately cylindrical, slightly divergent; length to width ratio: 2.7. Anterolateral accessory seta (I) and anterolateral seta (II) subequal in length, both setae inserted together distally at 5/6 length of caudal ramus, smooth. Posterolateral seta (III) missing. Outer terminal seta (IV) long and pinnate (length seta/length caudal ramus: 1.5), inserted subterminally and projecting outwards; inner terminal seta (V) without fracture plane. Terminal accessory seta (VI) short (length seta/length caudal ramus: 0.7) and smooth. Dorsal seta (VII) articulate, inserted distally at 5/6 length of the caudal ramus, smooth.

ROSTRUM ( Fig. 4 a–b). Small, not demarcated at base, almost reaching distal margin of first antennulary segment, ornamented with two dorsal sensilla.

A1 ( Fig. 4 a–c). Prehensile, eight-segmented pocket-knife type sensu Schminke (2010). First segment short with transversal row of spinules; second segment longest, with six setae, longest seta unipinnate; third segment with four distal bare setae; fourth segment reduced to a small sclerite with two short setae; fifth segment enlarged with inner round expansion carrying one short and two longer subequal setae, and distal tubercle with one seta and one large aesthetasc, reaching past end of eighth segment; sixth segment bare, partially fused to previous one; seventh segment bare, distal anterior corner protruding as curved apophysis ending in bilobate tip; eighth segment with seven setae and apical acrothek represented two

setae and long, slender aesthetasc. Armature formula: 1-[0], 2-[1 unipinnate + 5 bare], 3-[4 bare], 4-[2 bare], 5-[4 bare + ae], 6-[0], 7-[0], 8-[7 bare + (2 bare + ae)].

A2 ( Fig. 4d ). Coxa unarmed; allobasis with transverse row of spinules on inner margin. Exp represented by small segment merged with allobasis, with pinnate apical seta. Enp bearing two short subdistal unipinnate spines and five longer distal elements: two of them geniculated, one transformed with furcate tip, all elements unipinnate, with long spinules near their insertions.

MDB ( Fig. 4e ). Coxal gnathobase bare, cutting edge with apical teeth and short bipinnate seta. Onesegmented palp, with two smooth, apical setae of equal length.

MX 1 ( Fig. 4f ). Praecoxal arthrite with three apical curved robust spines apically denticled, one subdistal curved seta. Coxal endite long, with one apical seta. Basis cylindrical, with two distal bare setae. Enp and exp absent (fused to basis without trace).

MX 2 ( Fig. 4g ). Basis with two endites, proximal endite short, with one thin, bare seta; distal endite cylindrical, longer, armed apically with two subequal thin bare setae and one transformed, leaf-like

pinnate seta; proximal endopodal segment drawn into apical unipinnate claw; distal endopodal segment with two long setae of equal length.

MXP ( Fig. 4h ). Subchelate, composed of small and unarmed syncoxa, basis slim and elongate, unarmed, 1-segmented enp fused to the claw-like apical seta.

P1 ( Fig. 5 a–b). With smooth and small intercoxal sclerite; coxa bare. Basis large, armed with small seta and lamellar hook with rounded tip on the inner margin near the enp insertion. Exp three-segmented, slightly shorter than endopod; exp-1 with thin pinnate seta on outer distal corner; exp-2 shortest and unarmed, ornamented with row of spinules along the subdistal and distal margins; exp-3 with two geniculate unipinnate apical setae and two subapical unipinnate setae. Enp two-segmented; enp-1 as long as first two segments of corresponding exp, with two transversal rows of few spinules on outer margin. Enp-2 thinner and shorter than enp-1, with two spinules at ¼ of inner margin and spinules along distal margin, with long, geniculate unipinnate seta and shorter unipinnate seta apically.

P2 ( Fig. 5c ). With smooth intercoxal sclerite, twice as wide as long, with concave distal margin. Coxa bare. Basis unarmed, with row of five spinules on outer margin. Exp three-segmented, exp-1 longest, with row of two spinules at ¼ of outer margin, longitudinal row of three spinules proximal to strong distolateral unipinnate spine. Second and third segments of same length; exp-2 unarmed, with row of spinules on distolateral corner, exp-3 armed with subapical outer unipinnate spine, apical bipinnate seta and outer unipinnate spine, ornamented with subapical spinule and inner hyaline frill. Enp onesegmented, about half length of exp-1, represented by cylindrical segment, with apical seta about as long as segment and three short spinules.

P3 ( Fig. 5d ). Intercoxal sclerite small, trapezoidal, unornamented, with slightly concave distal margin. Coxa bare. Basis robust, with long, slender, smooth outer seta and transverse spinule row above, and distal pore. Enp reduced to short seta. Exp-1 of characteristic shape: inner margin with large knob at ½ length and basal thin and pointed tubercle fused to exp; outer margin with proximal and distal rows of respectively one and four spinules. Exp-2 fused with exp-1, without ornamentation, prolonged into long apophysis slightly bent inwards, with pointed tip. Distal thumb represented by thin and pointed sigmoid segment, reaching past apophysis.

P4 ( Fig. 5 f–h). Intercoxal sclerite smaller than in P1–P3, with concave, smooth distal margin. Coxa bare. Basis armed with single slender seta on outer margin; ornamented with row of spinules at base of outer seta; row of three spiniform processes aligned along inner margin, of increasing size from outermost to innermost, all slightly curved inwards. Exp three-segmented, slender, all segments approximately of the same length; exp-1 slightly curved inwards, with distolateral pinnate spine, transversal spinular row at midlength of outer margin, spinular row at distolateral spine insertion, spinular row along distal margin; exp-2 unarmed, with spinular row along distal margin; exp-3 armed with outer unipinnate spine with spinules near insertion and very long apical pinnate seta, spine length less than ¼ of seta length; ornamentation represented by row of apical spinules and hyaline frill on inner distal corner. Enp onesegmented, as long as first two segments of corresponding exp, represented by curved plate with pointed inner tip carrying at outer border two outgrowths, distal one being elongate lamella, with undulating (crenulate) margins, reaching past end of enp, proximal outgrowth short curved lamella ending in thin tip.

P5 ( Fig. 6 a–d). Fused to intercoxal sclerite; represented by two trapezoidal cuticular plates with long basipodal seta and small pore near insertion. Armature on free distal margin, from inner to outer: one spiniform seta, three bare setae, outermost very short, remaining two subequal.

P6 ( Fig. 6a ). Vestigial, fused into simple cuticular plate, unornamented and unarmed. Adult female

HABITUS ( Fig. 2b ). Cylindrical and slender, without any demarcation between prosome and urosome. Free pedigerous somites without any lateral or dorsal expansions, all connected by well-developed arthrodial membranes. Integument weakly sclerotized, without cuticular pits, ornamented with sensilla on all somites except preanal one. Cuticular windows on urosomites and cephalothorax not present. Body length, excluding caudal setae, from 280 to 312 μm, mean 304 μm (n = 6), ornamentation of cephalothorax, somites, pigmentation and absence of nauplius eye as in male, except genital and first urosomite fused into double-somite. Cephalotorax representing about 19% of the total body length. Genital double-somite ( Figs 6e , 8c ) without any trace of subdivision. Genital field ( Figs 6e , 8c ) broader than tall, occupying anterior ventral ¼ of genital double-somite; single genital aperture covered by fused vestigial sixth legs; median copulatory pore located medially at ¼ of somite. Anal operculum convex.

CAUDAL RAMI ( Figs 6 f–g, 8b). Shape, ornamentation and armature similar to those of male, length to width ratio: 3.5.

ROSTRUM, A2 AND ORAL APPENDAGES. As in male.

A1 ( Fig. 7c ). Seven-segmented, aesthetasc on fourth segment longer than in male, reaching beyond end of seventh segment. First segment bare. Apical acrothek represented by two setae of different length and slender aesthetasc. Armature formula: 1-[0], 2-[1 unipinnate + 4 bare], 3-[4 bare], 4-[2 bare + ae], 5-[0], 6-[0], 7-[7 bare + (2 bare + ae)].

P1 ( Fig. 7d ). Intercoxal sclerite, coxa, basis as in male but lamellar hook missing; exp and enp similar to those of male in shape, ornamentation and armature.

P2 ( Fig. 7e ). Intercoxal sclerite longer and narrower than in male, coxa narrower than in male, basis as in male, but with spinular row below enp insertion. Exp as in male. Enp similar in shape and ornamentation to that of male, but with two apical spinule instead of three.

P3 ( Fig. 7 f–g). Intercoxal sclerite small, with concave margin, bare. Coxa bare. Basis with outer seta and spinular row near seta insertion. Exp two-segmented: exp-1 as long as exp-2, with distolateral curved unipinnate spine, transversal row of spinules at ¼ and ¼ of outer margin, hyaline frill on inner distal corner; exp-2 with outer unipinnate spine and apical pinnate seta, spine length about ¼ of seta, hyaline frill on inner distal corner. Enp represented by thin and pointed segment, slightly shorter than half of corresponding exp-1, apically pinnate.

P4 ( Figs 7h , 8d ). Intercoxal sclerite, coxa and exp as in male. Basis bare. Enp represented by thin cylindrical segment, about ¼ length of corresponding exp-1, ending in short bipinnate seta.

P5 ( Fig. 6e ). Fused to intercoxal sclerite, represented by two cuticular plates shorter than in male, with same armature and without distal pore.

P6 ( Fig. 6e ). Vestigial, fused into simple cuticular plate, covering gonopore, unornamented and unarmed.

Variability

The ornamentation of male P3 exp-1 can vary, as one specimen from Vucco Ucciardo has the proximal and distal rows of respectively two and three spinules ( Fig. 5e ). Two males ( Fig. 3b, d ) and one female ( Fig. 7b ) from Grotta del Banco di ferro, one male ( Fig. 3e ) and one female ( Fig. 7a ) from Vucco Ucciardo carry seven setae on the caudal rami instead then 6. In one female from Vucco Ucciardo the enp P4 is fused with the apical spine ( Fig. 7i ).

Accompanying fauna

Grotta del Banco di ferro, 10 May 2017: Maraenobiotus sp.: pool 4; Elaphoidella sp.: pool 2.

Figures

Fig. 2. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov. from Grotta superiore di Sant’Angelo (NHMUK). a. ♂, habitus. b. ♀, habitus. Scale bar: 50 µm.

Fig. 3. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov., ♂ (NHMUK). a. Anal somite, anal operculum and caudal rami, dorsal view. b. Anal somite, anal operculum and caudal rami, dorsal view (variability). c. Anal somite, anal operculum and caudal rami, dorsal view. d. Anal somite, anal operculum and caudal rami, lateral view (variability). e. Anal somite, anal operculum and caudal rami, lateral view (variability). a: Grotta superiore di Sant’Angelo; b, d: Grotta del Banco di ferro; c, e: Grotta Vucco Ucciardo. Scale bar: 50 µm.

Fig. 4. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov., ♂ (NHMUK). a. Rostrum and A1, ventral view. b. Rostrum and A1, ventral view. c. A1, disarticulated (antennular segments marked with Roman numerals). d. A2. e. Mdb. f. Mx1. g. Mx2. h. Mxp. a, e–g: Grotta Vucco Ucciardo; b–d, h: Grotta superiore di Sant’Angelo. Scale bar: 50 µm.

Fig. 5. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov., ♂ (NHMUK). a. P1, coxa, basis and enp, inner view. b. P1, basis and exp, outer view. c. P2, posterior view. d. P3, anterior view. e. P3, posterior view (variability). f. P4, anterior view. g. P4, intercoxal sclerite, coxa, basis and enp, outer view. h. P4, posterior view. a, c, e, g: Grotta Vucco Ucciardo; b, h: Grotta del Banco di ferro; d, f: Grotta superiore di Sant’Angelo. Scale bar: 50 µm.

Fig. 6. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov. (NHMUK). a. ♂, P5, P6, first and second urosomites, ventral view. b. ♂, P5. c. ♂, P5 and first urosomite, ventral view. d. ♂, P5. e. ♀, first urosomite, P5, P6, genital double-somite and genital field, ventral view. f. ♀, anal somite, anal operculum and caudal rami, dorsal view. g. ♀, anal somite, anal operculum and caudal rami, lateral view. a–b, e, g: Grotta Vucco Ucciardo; c: Grotta del Banco di ferro; d, f: Grotta superiore di Sant’Angelo. Scale bar: 50 µm.

Fig. 7. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov., ♀ (NHMUK). a. Anal somite, anal operculum and caudal rami, dorsal view (variability). b. Anal somite, anal operculum and caudal rami, lateral view (variability). c. Rostrum, A1. d. P1, anterior view. e. P2, inner view. f. P3, posterior view. g. Coxa, basis, enp P3, inner view. h. P4, posterior view. i. Coxa, basis, enp P4, inner view (variability). a, c–i: Grotta Grotta Vucco Ucciardo; b: Grotta del Banco di ferro. Scale bar: 50 µm.

Fig. 8. Cottarellicaris sanctiangeli Bruno & Cottarelli sp. nov. (CIME). a. ♂, anal somite, anal operculum and caudal rami, ventro-lateral view. b. ♀, anal somite, anal operculum and caudal rami, lateral view. c. ♀, P5, P6, genital double-somite and genital field, ventral view (outermost seta, which is folded behind the P5 arrowed). d. ♀, enp P4. Grotta superiore di Sant’Angelo. Scale bar: 50 µm.