Pagurus carmineus,

Lima, Daniel, Tavares, Marcos & Jr, Joel Braga De Mendonça, 2019, Paguroids (Decapoda: Anomura: Diogenidae and Paguridae) of the remote oceanic Archipelago Trindade and Martin Vaz, off southeast Brazil, with new records, description of three new species and , Zootaxa 4694 (1), pp. 1-63: 40-49

publication ID

https://doi.org/10.11646/zootaxa.4694.1.1

publication LSID

lsid:zoobank.org:pub:A24900FC-0D4F-499E-B7BB-566F6B7D350A

persistent identifier

http://treatment.plazi.org/id/87748C32-F833-F435-CE89-FBB0FA16F53F

treatment provided by

Plazi

scientific name

Pagurus carmineus
status

sp. nov.

Pagurus carmineus  sp. nov.

( Figs. 20C, DView FIGURE 20, 21–25View FIGURE 21View FIGURE 22View FIGURE 23View FIGURE 24View FIGURE 25)

Trindade specimens. HOLOTYPE: male 1.2 mm ( MZUSP 36402), Brazil, Espírito Santo, Trindade Island, stn Praia do Lixo , 20°31’29.8”S, 29°19’43.9”W, J.B. Mendonça coll., 25.x.2014, SCUBA, 13.7 mGoogleMaps  . PARATYPES: 1 ovigerous female ( MZUSP 36204), idem, stn Ponta Norte , 20°29’18.7”S, 29°20’18.3”W, J.B. Mendonça coll., 23.vi.2016, SCUBA, 11.3 mGoogleMaps  ; 1 ovigerous female ( MZUSP 36207), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonça coll., 19.vi.2016, snorkeling, 1.0 mGoogleMaps  ; 4 females (3 ovigerous) ( MZUSP 36206), idem, Praia do Parcel , 20°31’11.4”S, 29°18’00.6”W, J.B. Mendonça coll., 15.vii.2012GoogleMaps  ; 1 male ( MZUSP 36220), idem ,, Ponta Norte , 20°29’18.7”S, 29°10’18.3”W, J.B. Mendonça coll., 23.vi.2016, SCUBA, 11.3 mGoogleMaps  ; 1 male, 1 female (dissected) ( MZUSP 36205), idem, Enseada dos Portugueses, stn Farol , 20°29’52.3”S, 29°19’15.6”W, J.B. Mendonça coll., 06.vii.2015, artificial reef substrate, 12.6 mGoogleMaps  .

Non type material. 1 male ( MZUSP 36270), Brazil, Espírito Santo, Trindade Island, Ponta Norte , 20°29’18.7”S, 29°20’18.3”W, J.B. Mendonça coll., 01.iv.2014GoogleMaps  ; 7 males, 1 ovigerous female ( MZUSP 36285), idem, Ponta Norte , 20°29’18.7”S, 29°20’18.3”W, J.B. Mendonça coll., 23.vi.2016, artificial reef substrate, 11.3 mmGoogleMaps  ; 1 female ( MZUSP 36245), idem, Praia das Orelhas , 20°29’40.2”S, 29°20’32.9”W, J.B. Mendonça coll., 21.vii.2015, SCUBA, 15 mGoogleMaps  ; 1 female ( MZUSP 36275), idem, Praia das Orelhas , 20°29’40.2”S, 29°20’32.9”W, J.B. Mendonça coll., 20.x.2014, SCUBA, 14.1 mGoogleMaps  ; 1 female ( MZUSP 36282), idem, Praia das Orelhas , 20°29’40.2”S, 29°30’32.9”W, J.B. Mendonça coll., 01.vii.2016, SCUBA, 9.2 mGoogleMaps  ; 1 female ( MZUSP 36314), idem, Praia das Orelhas , 20°29’40.2”S, 29°20’32.9”W, J.B. Mendonça coll., 20.x.2014GoogleMaps  ; 2 males, 2 females ( MZUSP 36244), idem, Enseada dos Portugueses, stn Farol , 20°29’52.3”S, 29°19’15.6”W, J.B. Mendonça coll., 06.vii.2015, artificial reef substrate, 12.6 mGoogleMaps  ; 1 male ( MZUSP 36247), idem, Enseada dos Portugueses, stn Farol , 20°29’52.3”S, 29°19’15.6”W, J.B. Mendonça coll., 08.v.2014, SCUBA, 13.2 mGoogleMaps  ; 1 male, 1 female ( MZUSP 36283), idem, Enseada dos Portugueses, SECON/ECIT, 20°30’20.9”S, 29°18’43.7”W, J.B. Mendonça coll., 04.vii.2016, SCUBA, 11.4 mGoogleMaps  ; 2 females ( MZUSP 36280), idem, Enseada dos Portugueses, stn Farol , 20°29’52.3”S, 29°19’15.6”W, J.B. Mendonça coll., 22.iv.2014, SCUBA, 13.7 mGoogleMaps  ; 4 males, 1 female ( MZUSP 36284), idem, Ponta do Monumento , 20°30’10.3”S, 29°30’36.1”W, J.B. Mendonça coll., 02.iv.2014, SCUBA, 8.4 mGoogleMaps  ; 1 male ( MZUSP 36322), idem, Ponta do Monumento , 20°30’10.3”S, 29°20’36.1”W, J.B. Mendonça coll., 02.iv.2014, SCUBA, 8.4 mGoogleMaps  ; 2 males ( MZUSP 36246), idem, Ponta da Calheta , 20°30’18.72”S, 29°18’31.67”W, J.B. Mendonça coll., 25.vi.2015, artificial reef substrateGoogleMaps  ; 1 male ( MZUSP 36377), idem, Ponta da Calheta , 20°30’18.7”S, 29°18’31.6”W, J.B. Mendonça coll., 20.v.2014, SCUBA, 16.3 mGoogleMaps  ; 2 males ( MZUSP 36273), idem, Ilha da Racha , 20°30’26.5”S, 29°20’48.0”W, J.B. Mendonça coll., 01.vii.2016, SCUBA, 21.4 mGoogleMaps  ; 1 male ( MZUSP 36243), idem, Ilha da Racha , 20°30’26.5”S, 29°20’48.0”W, J.B. Mendonça coll., 01.vii.2016, SCUBA, 21.4 mGoogleMaps  ; 1 ovigerous female ( MZUSP 36218), idem, Ilha da Racha , 20°30’26.5”S, 29°20’48.0”W, J.B. Mendonça coll., 01.vii.2016, SCUBA, 21.4 mGoogleMaps  ; 1 male ( MZUSP 36281), idem, Ponta dos Andradas, 20°30’45.7”S, 29°18’21.9”W, J.B. Mendonça coll., 21.vii.2013, tide poolGoogleMaps  ; 1 male ( MZUSP 36276), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonça coll., 19.vi.2016, intertidal and snorkeling, 1.0 mGoogleMaps  ; 1 male, 1 ovigerous female ( MZUSP 36378), idem, Parcel das Tartarugas, stn tide pool, 20°31’10.4”S, 29°17’58.4”W, J.B. Mendonça coll., 19.vi.2016, intertidal and snorkeling, 1.0 mGoogleMaps  ; 1 female ( MZUSP 36278), idem, Enseada da Cachoeira, Farrilhões , 20°31’22.4”S, 29°19’52.0”W, J.B. Mendonça coll., 04.vii.2012, SCUBA, 17.9 mGoogleMaps  ; 1 male, 1 female ( MZUSP 36274), idem, Praia do Lixo , 20°31’29.8”S, 29°19’43.9”W, J.B. Mendonça coll., 22.iv.2014, SCU- BA, 14.5 mGoogleMaps  ; 5 ovigerous females, 11 males ( MZUSP 36211), idem, Praia do Lixo , 20°31’29.8”S, 29°19’43.9”W, J.B. Mendonça coll., 25.x.2014, SCUBA, 13.7 mGoogleMaps  ; 2 ovigerous females, 1 female, 1 male ( MZUSP 36277), idem, Praia do Lixo , 20°31’29.8”S, 29°19’43.9”W, J.B. Mendonça coll., 25.x.2014, SCUBA, 13.7 mGoogleMaps  ; 1 male ( MZUSP 36323), idem, Enseada do Príncipe, stn Paredão , 20°31’22.4”S, 29°19’52”W, J.B. Mendonça coll., 09.vii.2013, SCUBA, 19.6 mGoogleMaps  .

Comparative material examined: Pagurus brevidactylus  : 1 male ( MZUSP 34613), United States, Texas, Murphy’s Reef off South Padre Island , 26°23’N, 97°00’W, M. Wicksten coll., 10.viii.2006GoogleMaps  , SCUBA, 32 m; 1 male, 1 ovigerous female ( MZUSP 36038), Brazil, Paraíba, João Pessoa, Naufrágio do “Queimado”, 07°05.070’S, 34°44.852’W, L.R. Simone coll., SCUBA, 15 mGoogleMaps  ; 1 male, 1 female ( MZUSP 9440), Brazil, Pernambuco, Fernando de Noronha, Ponta da Sapata , P.S. Young coll., 03.xii.1988  ; 1 male ( MZUSP 36225), Brazil, Pernambuco, Fernando de Noronha, Ilha da Rata, Buraco do Inferno , 03°48’31.24”S, 32°22’52.71”W, L.R. Simone coll., 05.v.2013GoogleMaps  , 5– 10 m; 1 male ( MZUSP 13380), Brazil, Bahia, Salvador, Praia de Itapuã , L.R. Simone coll., 23–27.ii.1997  ; 1 male ( MZUSP 22153), Brazil, Rio de Janeiro, Angra dos Reis, Ilha Jorge Grego , V.S. Amaral & A.P. Dornellos coll., 08/ 09.viii.2009  ; 1 male ( MZUSP 28040), Brazil, Bahia, Boipeba, Castelhanos , sta 8, 13°39.957’S, 38°53.582’O, Tavares et al. coll., 19.ix.2012GoogleMaps  ; 1 male, 1 female ( MZUSP 28300), Brazil, São Paulo, São Sebastião, Arquipélago de Alcatrazes, sta Ponto 1— Costa Sul Portinho , 24°06’22.01”S, 45°41’36.26”W, L.R. Simone et al., coll., 04.xii.2012GoogleMaps  ; 2 males (32772), Brazil, São Paulo, São Sebastião, Parcel da Pedra Lisa, sta Ponto 5, 23°46’S, 45°15’W, IV Workshop of Neograstropoda coll., 25–30.viii.2014GoogleMaps  , 0–10 m; 1 male ( MZUSP 13808), Brazil, Santa Catarina, Porto Belo, xii.1998  ; 1 male ( MZUSP 16961), Brazil, Santa Catarina, Ilha do Arvoredo, Andrea-FURG coll  .

Type locality. Praia do Lixo , 20°31’29.8”S, 29°19’43.9”W, 13.7 m, Trindade Island, off Espírito Santo, BrazilGoogleMaps  .

Distribution. So far know only from Trindade Island, Espírito Santo, Brazil, between the intertidal and 21 m.

Etymology. The specific epithet is formed by the Latin word “carmin” (bright red) and the Latin suffix “ineus” (having the state or condition of), in reference to the red carmin color of the ocular peduncles and lateral margins of the carapace.

Description. Ten pairs of biserial gills; third maxilliped with only 1 rudimentary gill ( Fig. 20C, DView FIGURE 20). Shield ( Fig. 21AView FIGURE 21) as long as broad or slightly broader than longer; dorsal surface with tufts of short setae near to anterior margin. Anterolateral margin sloping; anterior margin between rostrum and lateral projection concave. Posterior margin truncate. Rostrum broadly rounded, lacking terminal spine. Lateral projections roundly triangular. Posterior carapace membranous; posteromedian plate with 2 tufts of long setae; branchiostergite with few tufts of moderately long setae laterally.

Ocular peduncle ( Fig. 21AView FIGURE 21) short, stout, cylindrical, with 0.4 times as long as shield, slightly dilated basally, 2.2 times as longer than broad; cornea slightly dilated; dorsomesial margin with row of tufts of short to moderately long setae. Ocular acicles subquadrate; terminal margins developed into 3–4 spines projections, separated by about basal width of 1 acicle.

Antennular peduncle ( Fig. 21AView FIGURE 21) long, overreaching distal corneal margin and antennal peduncles. Ultimate segment with distal tuft of long setae on dorsolateral margin, and scattered short setae on dorsal surface. Penultimate segment glabrous. Basal segment with small spine on laterodistal margin.

Antennal peduncle ( Fig. 21AView FIGURE 21) overreaching distal corneal margin. Fifth segment with tuft of long setae on distal margin, and scattered short setae on dorsal and ventral surfaces. Fourth segment with tuft of short setae on lateral surface, and scattered short setae on dorsal and ventral surfaces. Third segment with spine and tuft of long setae on ventrodistal margin. Second segment with dorsolateral distal angle produced into strong, acute projection terminating in small simple spine and small spine subdistally on ventral margin; dorsomesial distal angle with small sharp spine. First segment with strong spine on ventrodistal margin and small spine on lateral surface. Antennal acicle reaching or slightly overreaching proximal corneal margin; curved outward, terminating in strong spine; tip with tuft of long setae. Flagellum long, reaching tip of extended right cheliped; each article with 3–4 short setae (to about one-half of 1 article in length).

Mouthparts ( Fig. 22View FIGURE 22). Mandible without distinguishing characters. Maxillule with proximal endite with distal margin rounded; endopodite with external lobe well-developed, not curved, glabrous; internal lobe reduced, with very long terminal setae. Maxilla with endopodite inflated proximally, narrowing distally. First maxilliped with endopodite short; basal segment of exopodite inflated proximally, narrowing distally. Second maxilliped with basisischium fusion complete. Third maxilliped with basis-ischium fusion incomplete; basis with 1 small spine; ischium with crista dentata well-developed (9–10 spines, 2 stiff setae), 1 accessory tooth. Sternite of third maxilliped broad, with median acute projection.

Chelipeds unequal, right longer and stronger than left. Right cheliped ( Fig. 23View FIGURE 23 A–C) with fingers leaving broad hiatus when closed, terminating in small corneous claw, with tufts of short setae on distal margins. Dactylus shorter than palm; dorsal surface with scattered blunt spines; mesial surface with row of tubercles; cutting edge with small calcareous teeth and prominent tooth proximally. Fixed finger dorsal surface with proximal row of minute spines; cutting edge with row of small calcareous teeth and rather prominent calcareous tooth about midlength of finger. Palm ovate, 0.8 times as long as carpus; dorsal and dorsomesial surfaces with scattered spines; dorsolateral margin delimited by row of blunt spines extending from about midlength of propodus to the proximal part of fixed finger. Carpus trapezoidal, 1.1 times as long as merus; dorsomesial and dorsolateral margins with tufts of dense long setae; dorsal surface with few transversal rows of long setae; dorsomesial margin with irregular row of spines; dorsolateral margin unarmed (with row of spines in females). Merus subtriangular, dorsodistal margin unarmed, with row of moderately long setae. Ischium unarmed; dorsal and ventral surfaces with scattered moderately long and long setae.

Left cheliped (24 A–E) fingers leaving narrow hiatus when closed, ventrally spoon-shaped terminating in small corneous claw. Dactylus 1.9 times as long as palm; unarmed; dorsal and mesial surfaces with tufts of long setae; cutting edge with small corneous teeth. Fixed finger cutting edge with small calcareous teeth interspersed with small corneous teeth. Palm 0.4 times as long as carpus; dorsal surface markedly elevated, 2 median rows of strong spines, diminishing in size distally, merging into 1 row extending onto fixed finger; dorsomesial margin unarmed ( Fig. 24A, BView FIGURE 24) or with scattered tubercles ( Fig. 24DView FIGURE 24); dorsolateral margin with row of small spines, diminishing in size distally; lateral surface with row of small tubercles and tufts of short setae. Carpus almost as long as merus; 2 strong dorsodistal spines each preceded by one small spine; ventrolateral margin with 0–1 strong spine; ventromesial margin unarmed. Merus subtriangular; with row of moderately long setae on dorsal surface; distal margin with row of moderately long setae; ventrolateral margin with 1–3 strong spines; ventromesial margin with strong spine. Ischium unarmed.

P2 and P3 ( Fig. 23D, EView FIGURE 23) similar from left to right, slightly overreaching chelipeds. P3 longer than P2. Dactyli as long as or slightly smaller than propodi; curved, terminating in sharp corneous claw; ventromesial margin with 4–5 corneous spinules, diminishing in size distally. Propodi 0.7–0.8 times as long as carpi; ventrodistal margin with 1–2 corneous spinules, preceded by 0–1 spinule. Carpi 0.7–0.8 times as long as meri; dorsodistal margin with strong spine. Meri laterally compressed; that of P2 often with small spine on ventrolateral margin. Ischia unarmed. Sternite XII ( Fig. 21CView FIGURE 21) bilobed, separated by thin membranous cleft; anterior lobe subsemicircular, with tuft of long setae on distal margin.

P4 semichelate ( Fig. 24FView FIGURE 24). Dactylus curved, subtriangular, terminating in sharp corneous claw, 9–10 minute corneous teeth on ventrolateral margin; preungual process absent. Propodal rasp consisting of 2–3 rows of corneous scales. Carpus and merus with tuft of long setae on dorsodistal margin and dorsal surface medially. Ischium with tuft of long setae on ventrodistal margin.

P5 chelate. Propodal rasp occupying nearly half of lateral surface of propodus. Sternite XIV ( Fig. 24DView FIGURE 24), subdivided into 2 nearly symmetrical lobes.

Uropods strongly asymmetrical, left largest. Telson nearly symmetrical, with distinct transverse identation; posterior lobes separated by moderately deep U-shaped median cleft; terminal margins oblique, each armed with 3–4 small spines; lateral margin smooth, with chitinous narrow plate ( Fig. 21BView FIGURE 21).

Males with unpaired Pl3–Pl5, each with exopodite well-developed, endopodite reduced. Females with unpaired Pl2–Pl5; Pl2–Pl4 with both rami well developed, Pl5 with exopodite well-developed, endopodite reduced. Ovigerous females carrying few, large eggs ( Fig. 25A, BView FIGURE 25).

Color. In life ( Fig. 25View FIGURE 25), anterolateral region of shield with clusters of carmin red dots on a whitish background; dots becoming lighter and spread posteriorly. Branchiostergitegal regions carmin red anteriorly, red fading posteriorly. Ocular peduncles carmin red, corneas lighter red. Ocular acicles predominantly white with minute, reddish dots. Antennular peduncles pale blue; antennular flagellum with bicolor setae, bright yellow priximally and dark blue medially and distally. First and second antennal segments carmin red; remaining antennal segments mostly translucents occasionally with one pale red-brown band each. Male cheliped whitish, with yellowish/brownish patches on the dorsolateral and dorsomesial mergins; meri with two dorsal, one lateral, and one mesial reddish stripes on a bluish background. Female chelipeds propodi yellowish, fixed and movable fingers whitish; carpi yellowish, brownish proximally.Ambulatory legs with red brown, short, longitudinal stripes on a bluish background on dorsal, ventral, lateral and mesial surfaces, in otherwise whitish segments; dactyli bluish medially and distally.

Remarks. Pagurus carmineus  sp. nov. is morphologically closely related to the “provenzanoi group” of species with multispinose ocular acicles, especially P. brevidactylus  . However, P. carmineus  sp. nov. can be readly differentiated from P. brevidactylus  in having the propodal rasp consisting of 2–3 rows of ovate corneus scales ( Fig. 24FView FIGURE 24), whereas the propodal rasp has 4–6 rows of ovate corneus scales in P. brevidactylus  . The new species can be further distinguished by the ratio length/width of the ocular peduncles, markedly short and stout, about twice as long than wide in P. carmineus  sp. nov. ( Fig. 21AView FIGURE 21), whereas the ocular peduncles are distinctly longer and slender, about four times as long than wide, in P. brevidactylus  .

True, the use of the ratio length/width of the ocular peduncles as a distinguishing character between species of paguroids should be used cautiously, as the ratio length/width varies over ontogeny and juveniles tend to have short- er and stouter ocular peduncles than adults. And in some species, the females can become ovigerous before attaining the ratio length/width of full-sized adults, e.g. Clibanarius tricolor  (see Provenzano 1960). However, in some species of micro-pagurids recently discovered the adults had stout eyes and diminutive size ( Lemaitre et al. 2017). This is also the case of P. carmineus  sp. nov., of which we have examined 42 males (largest sl 1.2 mm, smallest sl 0.4 mm) 15 females (largest sl 1.0 mm, smallest sl 0.7 mm) and 15 ovigerous females (largest sl 1.2 mm, smallest sl 0.7 mm). Commonly, largest juveniles and smallest adults of decapod crustaceans overlap over a size range at their transition ( DeMartini et al. 2005; González-Pisani et al. 2017). In P. carmineus  sp. nov. it seems that the shield length of the largest juvenile female and smallest adult female overlap between 0.7 mm and 1.0 mm; overlapping and full grown females (> 1.0 mm sl) do not vary in terms of ratio length/width of the ocular peduncles.

Pagurus carmineus  sp. nov. is distinctly smaller (sl minimum–maximum = 0.7–1.2 mm; sl mean ± SD = 0.9 ± 0.1 mm; N= 53 spec) than its congeners of the “provenzanoi group” of species with multispinose ocular acicles, such as P. brevidactylus  (sl minimum–maximum = 1.0–4.0 mm; sl mean ± SD = 2.0 ± 0.4 mm; N= 195 spec) ( Lima et al. 2014). The new species is striking for carrying few, large eggs (minimum–maximum eggs/spec = 2–22; eggs/ spec mean ± SD = 10.4± 6.5; egg minimum–maximum major diameter = 0.3–0.5 mm; egg major diameter mean ± SD = 0.40 ± 0.05 mm, Fig. 21A, BView FIGURE 21). Conversely, P. brevidactylus  carry many small eggs (minimum–maximum eggs/spec = 27–1075 eggs; eggs/spec mean ± SD = 158 ± 119; egg major diameter mean ± SD = 0.38± 0.04 mm) ( Iossi et al. 2005).

Komai & Osawa (2001: 1295, fig. 3H) reported that Pagurus decimbranchiae Komai & Osawa, 2001  , had only one arthrobranch on the third maxilliped greatly reduced to a single rudimentary bud, in lieu of having a pair of well developed biserial arthrobranchs. Komai & Osawa (2001) assigned P. decimbranchiae  to the Pagurus anachoretus Risso, 1827  , species group with which P. decimbranchiae  share a number of morphological characters. Hitherto, no other species of Pagurus  is known to have a rudimentary bud on the third maxilliped. The finding herein that P. carmineus  sp. nov. also have only one arthrobranch on the third maxilliped reduced to a non-lamelate bud ( Figs. 20C, DView FIGURE 20, 22AView FIGURE 22) prompted us to investigate the gill formula of other species of the “provenzanoi group” from the southwestern Atlantic (SWA). As a result, we have found that the arthrobranch on the third maxilliped is also reduced to a single bud in P. aff. brevidactylus  from Trindade ( Fig. 20A, BView FIGURE 20), P. brevidactylus  s. str., P. criniticornis  , P. leptonyx Forest & de Saint Laurent, 1968  , and P. provenzanoi  . A fifth species of the “provenzanoi group” also known from the SWA, P. trichocerus Forest & de Saint Laurent, 1968  , only known from the male holotype and most probably a synonym of P. leptonyx  , has not been examined. The “provenzanoi group” is amphi-American in distribution and consists of 26 species (12 western Atlantic and 14 eastern Pacific species).

Recent molecular analyses based only on a few species suggested the monophyly of the “provenzanoi group” ( Olguín & Mantelatto 2013; Noever & Glenner 2017). Although many more species from the “provenzanoi group” await investigation, whether by molecular or morphological techniques, the reduction of the arthrobranch on the third maxilliped to a single bud may be a promising synapomorphy defining morphologically this species group.

Kingdom

Animalia

Phylum

Arthropoda

Class

Malacostraca

Order

Decapoda

Family

Paguridae

Genus

Pagurus