Perinereis floridana ( Ehlers, 1868 ), 1100

Conde-Vela, Víctor Manuel, 2022, Reinstatement of Perinereis bairdii (Webster, 1884) and description of P. websteri sp. nov. from Bermuda, including the reproductive morphology of two Atlantic Perinereis species (Annelida: Errantia: Phyllodocida), European Journal of Taxonomy 787 (1), pp. 104-145 : 126-132

Perinereis floridana ( Ehlers, 1868)

Figs 1 , 10–11 , 13

Nereis floridana Ehlers, 1868: 503–506 .

Perinereis floridana – Langerhans 1880: 19 (new combination only). — Monro 1933c: 256. — Salazar- Vallejo & Jiménez-Cueto 1997: 367, figs 8, 32–33 (partim). — De León-González & Solís-Weiss 1998: 683–684, figs 6–7.

Type material examined

Syntypes UNITES STATES OF AMERICA • 11 ♂♂; Florida, Charlotte Harbor , Captiva Key ; G.W. Wurdemann leg.; MCZ ANNa-119 .

Additional material examined

UNITED STATES OF AMERICA • 1 spec.; Florida, Charlotte County, Lemon Bay ; 26°54′50″ N, 82°20′26″ W; 11 Jan. 1938; O. Hartman leg.; LACM-AHF 7377 • 3 specs; same collection data as for preceding; 14 Jan. 1938; O. Hartman leg.; LACM-AHF 7376 .

MEXICO • 16 specs; Yucatan, Isla Cerritos ; 18 Jan. 1991; Sergio I. Salazar-Vallejo leg.; ECOSUR P1100 • 1 spec.; Quintana Roo , Isla Contoy ; 11 Oct. 1993; Sergio I. Salazar-Vallejo leg.; ECOSUR P1112 .

Description

Epitoke

BODY AND MEASUREMENTS. Syntypes (MCZ ANNa-119) heteronereis males complete, in good conditions, one selected for description, 32 mm long, 3 mm wide at chaetiger 10 excluding parapodia, 110 chaetigers ( Fig. 10A ). Pigmentation not observed in all specimens ( Fig. 10A ).

PROSTOMIUM. Subpentagonal, 1.5 × as wide as long, anterior region distally entire, shorter than posterior region, dorsal groove present ( Fig. 10B ); anterolateral gap between antenna and palpophore 1.5× as long as diameter of antennae ( Fig. 10B ).

ANTENNAE. Cirriform, not passing palps, half as long as prostomium, gap between them as long as basal wide of antennae ( Fig. 10B ).

PALPS. Palpophores subconical, swollen, 1.6 × as wide as long, shorter than prostomium, subdistal transverse groove present ( Fig. 10B ). Palpostyles rounded.

EYES. Rounded, anterior and posterior pairs subequal, in rectangular arrangement, anterior and posterior pair overlapped, posterior pair not covered by anterior margin of tentacular belt ( Fig. 10B ).

TENTACULAR BELT. As long as chaetiger 1, anterior dorsal margin omega-shaped ( Fig. 10B ).

TENTACULAR CIRRI. Smooth, cirrophores conspicuous, longest cirri extending backwards up to end of chaetiger 7 ( Fig. 10B ).

PHARYNX. Everted ( Fig. 10B ); jaws brown, 7–8 teeth, proximal ones ensheathed ( Fig. 10C–D ). Maxillary ring: I = 2 cones in a vertical row; II = 6–7 cones in arc; III = 9 cones, 7 in a central ellipse and one cone at each lateral side of the ellipse; IV = 13–14 cones in arc ( Fig. 10C–D ). Oral ring: V = 1 cone displaced toward posterior margin of ring; VI = 1–1 smooth bar; VII–VIII = 24 cones in two bands: anterior band with one furrow row with 6 cones with 1 cone on the regions a–c, and 1 cone in ridge region A; second band with one furrow row with 8 cones with 1 cone on the regions a–d, and one ridge row with 9 cones, 3 cones in region A and 1 cone in each regions B–D ( Figs 10C–D , 13 ); rarely anterior and posterior bands very close to each other ( Fig. 10D ). Furrow pattern of areas VI–V–VI, λ-shaped ( Figs 10C , 13 ).

BODY REGIONS. Two regions: 1) pre-natatory region includes chaetigers 1–17, subdivided into two subregions: a) dorsal cirri in chaetigers 1–7 and ventral cirri in chaetigers 1–5 pyriform or cattail-like, and 2) chaetigers 8–17 with both dorsal and ventral cirri cirriform; 2) natatory region includes chaetigers 18 to end of body, chaetiger 18 with upper lamella in dorsal cirrus and both upper and lower lamellae in ventral cirrus, in chaetiger 19 and remaining ones also appear lower lamella in dorsal cirrus, a lamella below median ligule, ventral lamella and a basal, dorsal protrusion in neuropodial ventral ligules.

PRE- NATATORY REGION. Parapodia resembling atokous ones ( Fig. 10I–L ). Dorsal cirrus cattail-like in chaetigers 1–7, filiform thereafter ( Fig. 10I–L ); 1.5× as long as distal lobe of dorsal ligule in chaetigers 2–17 ( Fig. 10I–L ); 3.2 × as long as proximal lobe of dorsal ligule in chaetiger 2, 2.3× in chaetiger 6, 2× in chaetigers 10–17. Dorsal ligule digitiform in first chaetiger, becoming subconical with blunt tip toward posterior ones; distal lobe of dorsal ligule 1.3× as long as median ligule in chaetigers 6–10, 1.6× in chaetiger 17 ( Fig. 10I–L ). Median ligule subconical with blunt tip throughout; 3× as long as neuroacicular ligule in chaetiger 6, 3.5 × in chaetigers 10–17 ( Fig. 10I–L ). Neuroacicular ligule subconical with blunt tip throughout. Neuropodial superior and inferior lobes rounded, inferior one longer and wider than superior one throughout ( Fig. 10I–L ); neuropodial postchaetal lobe rounded, half as long as neuroacicular ligule. Ventral ligule digitiform throughout; 1.4 × as long as neuroacicular ligule in chaetiger 2, 1.8 × in chaetigers 6–10, 1.4× in chaetiger 17 ( Fig. 10I–L ). Ventral cirrus cattail-like in chaetigers 1–5, filiform thereafter; 0.7× length of ventral ligule throughout ( Fig. 10I–L ).

NATATORY REGION. Parapodia distinct from atokous ones ( Fig. 10M–O ). Dorsal cirrus filiform, crenulations present from chaetiger 18, decreasing in size and number toward posterior chaetigers until disappear ( Fig. 10M–O ); 1.7× as long as distal lobe of dorsal ligule in chaetiger 18, 1.4× in chaetiger 40, 1× in chaetiger 95 ( Fig. 10M–O ). Upper lamella of dorsal cirrus present since chaetiger 18, flabellate throughout, larger in middle chaetigers, as long as dorsal cirrus in chaetiger 40 ( Fig. 10M–O ); lower lamella since chaetiger 19, flabellate and much smaller than upper lamella of dorsal cirrus throughout ( Fig. 10M–O ). Dorsal ligule subconical with blunt tip in chaetiger 18, digitiform thereafter, becoming narrower toward posterior chaetigers ( Fig. 10M–O ); distal lobe of dorsal ligule 1.4× as long as median ligule in chaetigers 18–40, 1.2× in chaetiger 95 ( Fig. 10M–O ). Median ligule subconical with blunt tip in chaetiger 18, digitiform thereafter, becoming broader toward middle chaetigers ( Fig. 10M–O ); 4× as long as neuroacicular ligule in chaetiger 18, 2.6 × in chaetiger 40, 1.6× in chaetiger 95 ( Fig. 10M–O ); small basal lamella of median ligule present since chaetiger 19, flabellate throughout ( Fig. 10M–O ). Neuroacicular ligule subconical with blunt tip in chaetiger 18, digitiform thereafter, superior and inferior lobes absent ( Fig. 10M–O ). Neuropodial postchaetal lobe rounded in chaetiger 18, transformed into broad flabellate ventral lamella ( Fig. 10M–O ); 2× as long as neuroacicular ligule in chaetiger 40, 1.5× in chaetiger 95 ( Fig. 10M–O ). Ventral ligule subconical with blunt tip in chaetiger 18, digitiform with a small basal, dorsal lobe thereafter ( Fig. 10M–O ); 2 × as long as neuroacicular ligule in chaetiger 18, 1.2× in chaetiger 40, 1 × in chaetiger 95. Ventral cirrus filiform throughout; as long as ventral ligule throughout ( Fig. 10M–O ). Upper lamella of ventral cirrus divided in two subequal, digitiform lobes ( Fig. 10M–O ); lower lamella of ventral cirrus flabellate, 2–3× as wide as upper ones throughout ( Fig. 10M–O ).

ACICULAE. Basally amber and distally dark brown, amber region enlarges toward posterior chaetigers ( Fig. 10I–O ). Notoaciculae absent in first two chaetigers ( Fig. 10I ), proximal end rectangular in nonnatatory region and becoming flabellate in natatory one ( Fig. 10I–O ).

NOTO- AND NEUROCHAETAE. Resembling atokous ones in non-natatory region ( Fig. 10E–H ), replaced with paddle-like, heterogomph chaetae with short bosses in natatory region ( Fig. 10N–O ).

PYGIDIUM. Crenulated, with a rosette of papillae formed by two or three rows of papillae; anal cirrus as long as last 12–14 chaetigers ( Fig. 10A ).

Atoke

BODY AND MEASUREMENTS. Specimens from Isla Cerritos (ECOSUR P1100) in excellent condition, best preserved specimen complete, 52 mm long, 2.8 mm wide at chaetiger 10 excluding parapodia, 107 chaetigers. Pigmentation faded out; pale lines present, fingerprint-like pattern present from chaetiger 8.

PROSTOMIUM. Subtriangular, 1.2× as long as wide, anterior region distally entire, 1.7 × as long as posterior region, dorsal groove present ( Fig. 11A ); anterolateral gap between antenna and palpophore shorter than diameter of antennae ( Fig. 11A ).

ANTENNAE. Subconical, not passing palps, half as long as prostomium, gap between them shorter than basal wide of antennae ( Fig. 11A ).

PALPS. Palpophores subconical, swollen, 1.2× as long as wide, as long as prostomium, subdistal transverse groove present. Palpostyles rounded or digitiform.

EYES. Rounded, anterior and posterior pairs subequal, in trapezoidal arrangement, wide shorter than basal wide of antennae, sometimes posterior pair partly covered by anterior margin of tentacular belt ( Fig. 11A ).

TENTACULAR BELT. 1.4× as long as chaetiger 1, sometimes slightly covering posterior pair of eyes, anterior dorsal margin straight ( Fig. 11A ).

TENTACULAR CIRRI. Moniliform, not jointed, longest cirri reaching end of chaetigers 7–8 ( Fig. 11A ).

PHARYNX. Dissected; jaws brown, cutting edge with 7–8 teeth, proximal ones ensheathed ( Fig. 11H ). Maxillary ring: I = 2 cones in vertical line; II = 7–9 (7–10) cones in arc; III = 13 (10–14) cones in rectangle; IV = 16–17 (14–19) cones in arc. Oral ring: V = 1 cone; VI = 1–1 smooth bars; VII–VIII = 18 cones in two bands: anterior band with 4 paragnaths in a single furrow row with 1 cone on the regions a–b (sometimes 1 cone on regions c and A–B); posterior band with 6 cones in furrow row with 1 cone on the regions a–c (sometimes 1 cone on regions d), and 8 paragnaths in ridge row with 2 cones on region A and 1 cone on the regions B–D (sometimes 1 additional cone on region A) (Fig. 14). Furrow pattern of areas VI–V–VI, λ-shaped.

DORSAL CIRRI. Digitiform in first chaetigers, filiform thereafter, attached basally to dorsal ligule in anteriormost chaetigers, medially thereafter ( Fig. 11C–G ); 1.3× as long as distal lobe of dorsal ligule in chaetigers 2 and 10, 0.8–0.9× length thereafter; 3.7 × as long as proximal lobe of dorsal ligule in chaetiger 2, 2.3× in chaetiger 10, 0.8–0.9× length thereafter ( Fig. 11C–G ).

DORSAL LIGULES. Subconical with blunt tip in anterior and middle chaetigers, becoming pennant-like toward posterior chaetigers, with distal lobes longer than proximal ones in first chaetigers, becoming as long as in middle chaetigers and shorter than in posterior chaetigers ( Fig. 11C–G ). Distal lobe of dorsal ligule subconical throughout; 1.3× as long as median ligule in chaetiger 10, 1.8× in chaetigers 28 and 61, 2.4× in chaetiger 91 ( Fig. 11C–G ).

MEDIAN LIGULES. Digitiform in anterior chaetigers, becoming subconical with blunt tips in middle chaetigers and digitiform to lanceolate in posterior ones ( Fig. 11C–G ); 1.2 × as long as neuroacicular ligule in chaetiger 10, 1.8× in chaetigers 28 and 61, 1.5× in chaetiger 91 ( Fig. 11C–G ).

NEUROACICULAR LIGULES. Subconical with blunt tip throughout ( Fig. 11C–G ). Neuropodial superior and inferior lobes present in anterior chaetigers, both rounded, inferior one wider than superior one throughout ( Fig. 11C–G ).

NEUROPODIAL POSTCHAETAL LOBES. Rounded, half as long as neuroacicular ligule throughout.

VENTRAL LIGULES. Digitiform throughout ( Fig. 11C–G ); 1.4× as long as neuroacicular ligule in chaetiger 2, 1 × in chaetigers 10 and 28, 1.2× in chaetigers 61 and 91 ( Fig. 11C–G ).

VENTRAL CIRRI. Subconical throughout ( Fig. 11C–G ); 0.8× as long as ventral ligule in chaetiger 2, 0.5– 0.6× in chaetigers 10–61, 0.7× in chaetiger 91 ( Fig. 11C–G ).

NOTOCHAETAE. All homogomph symmetrical spinigers. Blades of spinigers with basal pectinate, coarse teeth, becoming minute toward distal end.

NEUROCHAETAE. Homogomph symmetrical spinigers and heterogomph falcigers in supra-acicular fascicles, heterogomph spinigers and falcigers in sub-acicular fascicles. Neuropodial homogomph spiniger as notopodial ones ( Fig. 11L ). Heterogomph spinigers with blades pectinate, minute teeth and decreasing in size toward distal end ( Fig. 11K ). Heterogomph falcigers pectinate, long and narrow teeth, 7/10 of inner edge of blade dentate, distal tips stout, stouter in supra-acicular falcigers ( Fig. 11I–J ).

PYGIDIUM. Crenulated, bilobate ( Fig. 11B ); anal cirri subulate, as long as last 5 chaetigers ( Fig. 11B ).

Remarks

Records of Perinereis floridana in the Gran Caribbean region are scarce ( Salazar-Vallejo 1996; Dean 2012). This species was described based on epitoke specimens, but most of its records are based on atokes. It is relevant because Perinereis species lack a caudal region with non-transformed parapodia, so parapodial features attributed to the atokes of P. floridana in posterior chaetigers do not come from the original description. Ehlers (1868) placed P. floridana in a group characterized by dorsal ligules of similar size along the body and not expanded or widened (in posterior chaetigers), parapodia with a similar development along the body, paragnaths in all pharyngeal areas, and with both conical and transverse (bars) paragnaths. The parapodial features can differ in atoke and epitoke specimens in Perinereis species as shown in this work, so possibly Ehlers justified the inclusion of P. floridana in this group using the paragnath arrangement and its comparison with another close species, P. cultrifera , which match with the mentioned features.

Perhaps the atoke morphology of P. floridana was also taken from Fauvel (1923) when he synonymized Nereis floridana and other 8 species with Perinereis cultrifera because of the presence of a single smooth bar on each area VI, although Fauvel recognized they differ in the number of paragnaths in the area V. Fauvel (1923) described parapodia of P. cultrifera as follows: dorsal and median ligules subequal, with dorsal cirri slightly surpassing tips of dorsal ligules, and both neuroacicular and ventral ligules shorter than median ligules, with short ventral cirri, in posterior parapodia the ligules become sharper and the notopodial ligules are longer than neuropodial ones. For example, Wesenberg-Lund (1949) recorded ‘ P. cultrifera Gr. var. floridana Ehl. ’ from Faror, Iran citing Fauvel and his paragnath-based grouping.

Fauvel’s synonymy was not followed in all cases. Horst (1924) recorded Nereis (Perinereis) floridana from Balabalagan islands and declared he followed Gravier instead. Hartman (1938a) examined the type material of P. floridana but omitted further comments because there was a redescription by Langerhans (1880: pl. 15 fig. 24a–e), but actually Langerhans (1880) described some atoke specimens from Madeira as Perinereis floridana based on their supposed similarity only without further comments. Monro (1933c) disagreed with Fauvel about the synonymy with P. cultrifera and reported P. floridana for Dry Tortugas, Florida, but without description or figures. Later, Hartman (1951) reported atokes and epitokes of P. floridana from Lemon Bay, Florida, and cited Monro (1933c), but some specimens differ in the following features: 1) area V has 3 paragnaths and 2) sometimes two bars in areas VI. Andrew & Andrew (1953) reported P. floridana from North Bimini and highlighted the species has middle chaetigers resembling posterior ones, contrary to P. anderssoni . Finally, Salazar-Vallejo & Jiménez-Cueto (1997) described atokes as P. floridana with specimens from the Yucatan Peninsula, and de León-González & Solís-Weiss (1998) described atokes and female epitokes using specimens from Florida and the Yucatan Peninsula, and they agreed on that P. floridana have dorsal cirri not extending beyond the tip of distal lobes of dorsal ligules in middle and posterior chaetigers, and dorsal ligules in posterior chaetigers slightly longer than in anterior and middle chaetigers but not widened.

Here, the set of features used to determine atokes specimens belong to P. floridana are from the tentacular belt, the pharynx, and the chaetigers (anterior chaetigers of atokes, chaetigers 8–17 in the pre-natatory region of epitokes): 1) length of longest tentacular cirri, 2) shape and number of teeth in jaws, 3) similar number and arrangement of paragnaths in oral ring, especially areas I and III, 2) position of single paragnath in area V and size of smooth bars in areas VI, 3) similar number and arrangement of paragnaths in areas VII–VIII, 4) similar neuroacicular falcigers in anterior chaetigers, 5) similar ratios among notopodial processes and between ventral ligules and ventral cirri in anterior chaetigers. Main parapodial differences (excepting dorsal and ventral cirri) between atokes (anterior chaetigers) and epitokes (non-natatory region) described here are 1) both notopodial and neuropodial ligules more rounded in epitokes, and 2) relative size of neuropodial ligules. There are similar differences between atokes and epitokes of P. bairdii and P. cariboea , which are discussed below.

Based on the main morphological atokous of P. floridana , i.e., the long tentacular cirri, the presence of a single cone in areas V and a single smooth bar on each area VI, the two rows of paragnaths in areas VII–VIII, and parapodia with slight changes between middle and posterior chaetigers, closely resembles P. websteri sp. nov., but they differ as discussed in the remarks section of the latter species. Santos (2007) recorded P. floridana from Rocas Atoll, Brazil, but the specimens described differ from P. floridana as follows: 1) in Brazilian specimens, chaetiger 1 has dorsal cirri 2.4× as long as the distal lobe of the dorsal ligule, whereas in P. floridana they are 1.3× longer; 2) in Brazilian specimens, middle and posterior chaetigers, have dorsal cirri longer than distal lobes of dorsal ligules, whereas in P. floridana they are subequal or shorter; 3) in Brazilian specimens, middle chaetigers have both dorsal and median ligules subequal, and in posterior chaetigers dorsal ligules are 1.5× as long as median ones, whereas in P. floridana dorsal ligules are 1.8 × as long as median ligules in middle chaetigers and 2.4× longer in posterior ones. Other records of P. floridana outside the Gulf of Mexico and the Caribbean Sea, as in the Indian Ocean (e.g., Horst 1924; Yousefi et al. 2011), are likely distinct species and need to be reassessed.

Regardless of the different prostomial features, the length of largest tentacular cirri, and the pharyngeal arrangement, males of P. floridana also differs from males of P. bairdii in the following key parapodial features: 1) in P. bairdii , the natatory region starts in chaetiger 14, whereas in P. floridana it starts in chaetiger 18; 2) in P. bairdii , the first chaetiger from the natatory region (14) has dorsal ligules larger than in the corresponding chaetiger in P. floridana (18); 3) in P. bairdii , both dorsal and ventral lamellae are shorter along the natatory chaetigers than in P. floridana ; 4) in P. bairdii , the median ligules along the natatory chaetigers are shorter than in P. floridana .

Distribution

Gulf of Mexico, Caribbean Sea.

Figures

Fig. 1. Parapodial morphology of species of Perinereis Kinberg, 1865 and points used to obtain segments and length ratios of parapodial structures described in this work. Arrows and labels correspond to points used for the measuring parapodial structures as described in Material and methods section. A. Parapodium from posterior chaetiger of Perinereis floridana (Ehlers, 1868) (ECOSUR P1100). B. Parapodium from natatory region of Perinereis floridana (LACM-AHF 7376). Abbreviations: Dc = dorsal cirrus; DDoL = distal lobe of dorsal ligule; DoL = dorsal ligule; LDc = lower lamella of dorsal cirrus; LoM = lower lamella of median ligule; LVc = lower lamella of ventral cirrus; MeL = median ligule; NeL= neuroacicular ligule; PDoL = proximal lobe of dorsal ligule; PoL = neuropodial postchaetal lobe; PreL = notopodial prechaetal lobe; UDc = upper lamella of dorsal cirrus; UVc = upper lamella of ventral cirrus; Vc = ventral cirrus; VeL = ventral ligule; VeLa = ventral lamella.

Fig. 10. Perinereis floridana (Ehlers, 1868). Syntypes of Nereis floridana (MCZ ANNa-119). A. Whole specimen, dorsal view. B. Anterior end, dorsal view. C. Pharynx, dorsal view. D. Pharynx, ventral view. E. Supra-acicular heterogomph falciger, chaetiger 6. F. Supra-acicular heterogomph falciger, chaetiger 17. G. Sub-acicular heterogomph falciger, chaetiger 6. H. Sub-acicular heterogomph falciger, chaetiger 17. I. Chaetiger 2, right parapodium. J. Chaetiger 6, right parapodium. K. Chaetiger 10, right parapodium. L. Chaetiger 17, right parapodium. M. Chaetiger 18, right parapodium. N. Chaetiger 40, right parapodium.O. Chaetiger 95, right parapodium. Scale bars: A = 2 mm; B = 1 mm; C–D = 0.5 mm; E–H = 10 μm; I–O = 0.2 mm.

Fig. 11. Perinereis floridana (Ehlers, 1868). Non-types (ECOSUR P1100). A. Anterior end, dorsal view. B. Posterior end, dorsal view. C. Chaetiger 2, right parapodium. D. Chaetiger 10, right parapodium. E. Chaetiger 28, right parapodium. F. Chaetiger 61, right parapodium. G. Chaetiger 91, right parapodium. H. Left jaw, dorsal view. I. Supra-acicular heterogomph falciger, chaetiger 61. J. Subacicular heterogomph falciger, chaetiger 61. K. Sub-acicular heterogomph spiniger, chaetiger 61. L. Supra-acicular homogomph spiniger, chaetiger 61. Scale bars: A–B = 1 mm; C–H = 0.25 mm; I–J = 10 μm; K–L = 50 μm.

Fig. 13. Arrangement of paragnaths in the ridge and furrow regions of areas VII–VIII in species of Perinereis Kinberg, 1865. Filled figures with solid edges represent paragnaths observed in all specimens examined, no-filled figures with dashed edges represent paragnaths observed in some specimens only.