Phoronidea from Brazil

Phoronidea from Brazil

Liliana Forneris

SUMMARY

1. The demarcation of the species of Phoronidea is difficult because of the great variability of the characters which does not enable the fixing of good differential characters yet. The studies of populations and of life-histories must be undertaken to allow an accurate systematic definition. Within the family Phoronidae, generally considered as composed of only one genus, Phoronis, groups of very divergent species can be distinguished, which must perhaps be considered as genera or at least sub-genera. Owing to the inter gradation of the specific characters within such groups, it is not possible to know if they really represent species of form minor specific units.

I. Phoronis hippocrepia Wright 1856 - from Santos, Cananéia and Ubatuba; in the first locality boring in oyster shells; in the second non-boring specimens; and in the third only larvae.

a) Phoronis (Figs. 1-33) - The single nephrostome is the main characteristic of the specimens; through the interruption of the lateral mesentery at the level of the nephrostome the communication between the anal and oral cavities is assured. Longitudinal muscle bundles 26-30 in number, arranged according to the formula: ; marginal fibres, three to five in number, on each side of the bundle. Giant fibres of variable diameter: 4.7 to 7.0 microns, the right one, and 3.6 to 10.0 microns, the left. Blood corpuscles from 6.4 to 9.6 microns in diameter. Lophophoral organs also present in specimens with embryos in the tentacular crown. Simultaneous hermaphrodite, at least the specimens from Santos. Eggs: 100 microns. Spermatozoa: 20 microns, with vesicular head. Seven to eight pairs of chromosomes.

b) Actinotrocha (Figs. 34-55 )- Primordium of metasoma appears in the larva with one pair of tentacles; peri-anal ciliated ring is present in the larva with four tentacles in which the anlage of the blood-corpuscle mass appears. Ganglion already developed in this larva. Metasoma of the four-tentacle larva forms a small invagination. Nephridia are not terminal any more. Rare glands present in the hood epithelium. In the larva with six tentacles the metasoma forms an ample sac; the primordium of the lophophore coelom appears. Stomach diverticulum absent or single without vacuolization. Nephridia open laterally to the metasoma aperture. In the advanced larva (eight to 10 tentacles) the metasoma forms a series of folds; its longitudinal muscles are differentiated into bundles; a large ventral mass of blood corpuscles is present ventral to the eosophagus and a few corpuscles are grouped on each side at the insertion level of the ventral tentacles. Lophophore coelom well developed; dorsal vessel present; a pair of ventral muscles in the trunk. On the dorsal surface of the hood there is an accumulation of gland cells perhaps the sensory organ. During metamorphosis the evaginated metasoma shows longitudinal muscle bundles. Larval tentacles degenerate. Afferent and efferent vessels are already formed.

c) The following observations were made: Maximum survival in aquarium: one year; observed breeding season: July to March; duration of individual reproduction: over 15 days; development: egg to young larva - four days; pelagic perid - 9 to 12 days; metamorphosis - two hours. Irregular blood circulation, with about 10 pulsations per minute (temperature of 24ºC). Food: mostly diatoms, also protozoans and detritus. Not necessarily a boring species. The occurrence in aggregations seems accidental. Maximum of 12 embryos in the brood-pouch. The larvae become free with four tentacles and, exceptionally, in aquarium, shortly before metamorphosis. Swim in more or less large circles, diving often; shortly before metamorphosis, swim on the bottom. Metamorphosis was obtained even without special substratum. Eurythermic species (temperatures from 16 to 27ºC) endures salinities from 30.63 to 37.38 ‰.

II. Phoronis ovalis Wright 1856 (Figs. 56-63) - from Santos, in oyster shells. A single median lophophoral organ (Figs. 62, l; 63, b) ; three to four marginal fibres along each side of the longitudinal muscle bundle; two giant fibres 2.5 microns in diameter (Fig. 61, k), not always noticed. Lower lip of the nephridial funnel short or long, prolonged downwards. Larva observed in March.

III. a) Actinotrocha bella (new technical name) - from plankton collected off the northeast coast of Brazil.

Stage with 18 to 22 tentacles (Fig. 64 A, B) - Metasoma present as a small tube; no mass of blood corpuscles, sensory organ and dorsal vessel. Peri-anal ciliated ring developed. A pair of retractors in the hood; a single diverticulum, vacuolated; nephridia placed above the septum, open laterally the aperture of the metasoma.

Stage with 24 tentacles (Fig. 71 A) - Definitive tentacles absent. Stomach diverticulum single. Metasoma represented by a small invagination. Nephridium constituted by a large nephridial canal. Subepithelial muscle layer present. No mass of blood corpuscles and sensory organ. Narrow body cavity.

Stage with 38 tentacles (Fig. 71, B, B₁) - Definitive tentacles absent. A large blood sinus (Fig. 76, d) ventral to the oesophagus, extending laterally to the mouth; on the lower side it stretches out along the ventral part of the diverticulum (Fig. 77, d) ending near the nephridial canal (Fig. 78, h). Sensory organ (Fig. 83, c) consists of accumulations of basophilous gland cells and a nerve deeply placed. Dorsal vessel (Fig. 79, p) and caecal vessels (l) developed. Metasoma with longitudinal muscles differentiated into bundles (Fig. 79, n). A pair of dorsal retractors in the hood (Figs. 76, 77, b). Two pairs of ventral muscles in the trunk. Peri-anal ciliated ring developed and undulated (Fig. 81). These advanced larvae were related to the young larvae mentioned above by similarities of structure. Only their rearing will enable to confirm they belong to the same ontogenetic sequence.

b) Actinotrocha chata n. t. n. (Fig. 90) - This young larva with 16 tentacles was found in plankton from Ubatuba. The metasoma presents itself in the stage of an epithelial thickening; no mass of blood corpuscles and sensory organ; ganglion present. It is probable that this larva represents a young A. wilsoni B with delayed development.

c) Actinotrocha wilsoni B - Larva distributed in the coastal waters of Ubatuba (100 km to the NE of Santos) towards the South down to lat. 28º00'S, off Lagoa Mangueira (State of Rio Grande do Sul) ; it is also found in the inlet waters of Cananéia and Paranaguá. Occurs July to February, maximum in October.

Euryhaline; it was found at temperatures from 18.5 about 27.0ºC. Larva with eight to 10 tentacles has no metasoma, no diverticulum, no mass of blood corpuscles; terminal nephridia open above the anus. In the 12-tentacle stage the metasoma appears as a thickening. In the stage with 16 to 18 tentacles the ventral pair of blood corpuscle masses appears. Nephridia then open laterally to the metasoma orifice. Metasoma represented by a small tube. The dorsal pair of blood corpuscle masses appears in the larva with 22 tentacles. Usually the definitive tentacles appear in the 22-tentacle larva. They reach a maximum of 26, the maximum of definitive ones being 20. In the mature larva the sensory organ is present (Fig. 118). Stomach diverticulum may be absent, paired or single, rarely vacuolated. Longitudinal muscles of the metasoma are differentiated into bundles. Anomalous metamorphosis may occur in the plankton. Larval tentacles are cast off shortly before metamorphosis. The pairs of blood corpuscle masses conjoin during metamorphosis. The evaginated metasoma showed 48 longitudinal muscle bundles and one nerve. The developed larva is transparent, with orange pigment; it swims on the bottom of the recipient in large circles, resting sometimes in an inverted position; the tentacles are a swimming aid, standing aloof from the body. The developed larvae are identical with Actinotrocha wilsoni B, whose young stages are not known. The present young stages described from plankton, were related to A. ivilsoni B, but only their rearing will confirm that they represent the ontogenetic stages of this larva.

RESUMO

Phoronis hippocrepia Wright de Santos, Cananéia e Ubatuba, vive no eulitoral, na zona de máxima vasante, perfurando conchas de ostra ou sobre fauna que recobre rochas. Hermafrodita simultânea, com fecundação precoce interna. A eliminação dos corpúsculos polares dá-se na câmara incubadora. Larvas aí se desenvolvem até o estádio de 4 tentáculos, excepcionalmente até o de 8 tentáculos. As seguintes observações foram feitas: máximo de sobrevivência em aquário: 1 ano; época de reprodução: julho a março; duração da reprodução individual: mais de 15 dias; desenvolvimento: do ôvo à larva jovem - 4 dias; em natação livre - 9 a 12 dias; metamorfose - 2 horas. Máximo de 12 embriões na câmara incubadora. O adulto, quase imóvel, é muito sensível quando recém-coletado; perde parte da sensibilidade quando no aquário. Circulação sanguínea irregular, com cerca de 10 pulsações por minuto. Alimentação: principalmente diatomáceas; também protozoários. Espécie não necessariamente cavadora. A ocorrência em agregados parece acidental. Larva nada em círculos, subindo à superfície e mergulhando em seguida; perto da metamorfose, nada no fundo. Espécie euritérmica (temperaturas de 16 a 27ºC), suporta salinidades de 30,63 a 37,38 ‰.

Phoronis ovalis Wright observada em Santos no mesmo biótopo de P. hippocrepia. As descrições anteriores acrescenta-se: órgão lofoforal mediano, Impar; fibras gigantes pares, 2,5 micra de diâmetro, nem sempre conspícuas; fibras marginais e centrais presentes em cada feixe muscular longitudinal; nefróstoma pequeno com sua parede inferior curta ou prolongada para baixo. A larva aberrante foi reconhecida dentro de pequenos tubos chitinosos, em março.

Actinotrocha bella (nome técnico novo) do plancton coletado ao largo da cidade de São Luís (E. do Maranhão) mostra, no máximo estádio obtido, 30 tentáculos larvais; os tentáculos definitivos são ausentes. Grande seio sanguíneo anterior. Divertículo impar do estômago. Metasoma desenvolvido, com musculatura longitudinal diferenciada em feixes. Órgão sensorial presente, constituído por uma acumulação de células glandulares basófilas. Retratores no capuz e no tronco. Vaso dorsal e lateral bem como capilares desenvolvidos. Camada muscular subepitelial presente.

Actinotrocha chata (nome técnico novo) do plancton de Ubatuba, refere-se a uma larva jovem com 16 tentáculos, que pode representar uma A. wilsoni B com desenvolvimento retardado.

Actinotrocha wilsoni B (nome técnico) das águas costeiras de Ubatuba para o sul até 34º41,5'S (altura da Lagoa Mangueira), e águas interiores de Cananéia e Paranaguá. No máximo estádio de desenvolvimento possui 26 tentáculos larvais e até 20 definitivos; estes dispõem-se em duas séries, uma de cada lado, sob os larvais. A larva é caracterizada pelas quatro massas de glóbulos sanguíneos que se fundem na metamorfose. Próximo da metamorfose os tentáculos larvais degeneram, permanecendo apenas os definitivos. O capuz começa a desintegrar-se 15 minutos após o início da evaginação do metasoma. Metamorfose no plancton ocorre, mas de modo anormal. Larva encontrada de julho a fevereiro com máximo em outubro. O adulto desconhecido, possui 48 feixes musculares longitudinais e uma única fibra gigante.

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7. ACKNOWLEDGMENTS

I wish here to express my deepest gratitude and appreciation to Professor Dr. E. Marcus, under whose supervision this work was done. Thanks are extended to Prof. Dr. W. Besnard, to Dr. J. de Paiva Carvalho and to Dr. Marta Vannucci for suggestions and criticisms. For the determination of part of the accompanying fauna I am much indebted to Mrs. Eveline du Bois-Reymond Marcus, Dr. Marta Vannucci, Prof. Dr. E. Marcus, Prof. F. C. Müller-Melchers, Dr. E. Nonato and Dr. J. de Paiva Carvalho. Dr. Marta Vannacci collected the first specimens and she and Dr. Tagea K. S. Bjornberg have kindly picked some Actinotrochae from plankton samples. The salinities were determined by the Division of Physical Oceanography (Dr. I. Emilsson).

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Publication Dates

• Publication in this collection
  15 June 2012
• Date of issue
  1959