Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

Two new species of Protocirrineris

(Polychaeta: Cirratulidae) from Brazil

Rodolfo Elías1, María A. Saracho-Bottero1 & Wagner F. Magalhães2

1. Instituto de Investigaciones Marinas y Costeras (IIMyC) - Departamento de Ciencias Marinas, Universidad Nacional

de Mar del Plata. Dean Funes 3350, B7602 AYL, Mar del Plata, Argentina, roelias@mdp.edu.ar,

asaracho@mdp.edu.ar

2. Water Resources Research Center, University of Hawaii at Manoa, Honolulu, Hawaii, USA, wagnerfm@hawaii.edu

Received 15-XI-2018. Corrected 06-II-2019. Accepted 30-VI-2019.

ABSTRACT. Introduction: Brazil has more than 8 000 km of coastline but its marine invertebrates are still

poorly known. The cirratulid polychaetes have been an especially neglected group because of its challenging

taxonomy. Objective: to describe a new species of Protocirrineris. Methods: In a recent survey in two of the

largest Brazilian bays, Todos os Santos Bay and Camamu Bay, 84 specimens of four morphotypes of the mul-

titentaculate genus Protocirrineris were collected. Because of the lack of modified chaetae, the taxonomy of

Protocirrineris is challenging and only two of these morphotypes are herein described as new species. Results:

These two new species are characterized by having tentacular filaments in two groups over chaetigers 3-4 and

first branchiae present from chaetiger 1. They differ from each other in relation to the nature of the achaetigerous

region, posterior region, ultrastructure of capillary chaetae and methyl green staining pattern. Two incomplete

specimens are also considered as Protocirrineris, and brief descriptions are given. Conclusions: The genus

Protocirrineris is cited for the first time in Brazil. Four new morphotypes were found in two large estuaries of

Brazil, and two new species are formally described.

Key words: Taxonomy, Protocirrineris, new species, Todos os Santos Bay, Camamu Bay, Northern Brazil.

Brazil has a coastline of more than 8 000

km and an extensive continental shelf. Although

the country has a large number of groups study-

ing polychaetes (Lana Da Cunha et al., 2017)

some families are still under-represented and

in need of taxonomic revisions. One of these

families is the Cirratulidae Ryckholt, 1851, in

which most reported species in Brazil represent

widely distributed taxa with low taxonomic

certainty. There is a general consensus that the

diversity of cirratulids is largely unknown and

underestimated and the largely used “cosmo-

politan” names are being erroneously applied

to local species (Tewary, 2015; Hutchings &

Kupriyanova, 2018; Elías, Saracho-Bottero, &

Simon, 2019 in this volume).

The first revision of the multitentaculate

cirratulids in Brazil applying recent taxonomic

standards was by Magalhães, Seixas, Paiva, &

Elias (2014), who recognized seven species

and a species complex. The two species widely

reported in the Brazilian grey literature, Tima-

rete filigera (Delle Chiaje, 1828) and Timarete

tentaculata (Montagu, 1808), were found to be

absent in shallow waters along the Brazilian

coast and probably have distributions limited

to their type localities in the Mediterranean and

North Atlantic, respectively (Magalhães et al.,

Elías, R., Saracho-Bottero, M. A., & Magalhães, W. F. (2019) Two new species of Protocirrineris

(Polychaeta: Cirratulidae) from Brazil. Revista de Biología Tropical, 67(S5) Suplemento,

S81-S91.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

2014). Other species include the species com-

plex Timarete punctata (Grube, 1859) that was

revealed to be composed of two distinct species

based on molecular data (Seixas, Zanol, Mag-

alhães, & Paiva, 2017), T. oculata (Treadwell,

1932), T. caribous (Grube, 1859) and T. ceci-

liae Magalhães, Seixas, Paiva, & Elias, 2014

together with Cirriformia chicoi Magalhães

et al., 2014 and C. capixabensis Magalhães

et al., 2014. The genus Protocirrineris Czer-

niavsky, 1881 has not been recorded in Brazil

up until now.

In recent surveys carried out in Todos os

Santos Bay (12.8° S - 38.7° W) and Camamu

Bay (13.9° S - 39° W) in northeastern Brazil,

four morphotypes of multitentaculate cirratu-

lids lacking specialized spines have been iden-

tified as Protocirrineris. The material turned

out to be composed of two undescribed species

described herein as Protocirrineris baiana sp.

nov. and Protocirrineris camamuensis sp. nov.

Two additional morphologically distinct spe-

cies were not formally named because only

incomplete specimens were collected. All the

diagnostic characters of Protocirrineris species

are summarized (Table 1).

MATERIAL AND METHODS

Eighty-four specimens of multitentaculate

cirratulids were collected at the lower and

median estuarine region of Paraguaçu estuary,

Todos os Santos Bay, northern Brazil (Mag-

alhães & Barros, 2011). These samples were

collected with a 0.3 m2 van Veen grab, washed

in the field with sea water through a 0.5 mm

mesh, preserved with 70 % ethanol (see Barros

et al., 2008 and Magalhães & Barros, 2011 for

more details). Additional cirratulid specimens

came from a collection in subtidal areas at

Camamu Bay, in shallow estuarine habitats

(Hatje et al., 2008; Pedreira, Barros, Farias,

Wagener, & Hatje, 2017).

All specimens were examined under a

dissecting scope, stereomicroscope, and light

microscope while selected individuals were

observed under a Scanning Electron Micro-

scope (SEM). Material was processed by a

JEOL, JSM-6460 LV scanning electron micro-

scope in the Mar del Plata National Uni-

versity. The material for SEM was prepared

performing dehydration in ethyl alcohol (70,

80, 90, 95 and 100 %). Samples were dried in

HMDS (hexamethyldisilazane), mounted on

aluminum stubs and sputter coated with Gold

Palladium. Type and voucher specimens are

deposited at the Museu de Zoologia da Uni-

versidade de São Paulo, São Paulo (MZUSP),

Brazil. Zoobank link: LSIDurn:lsid:zoobank.

org:pub:E03A2C2E-4102-4941-A2EF-

50CD49742771

RESULTS

Family Cirratulidae Ryckholt, 1851

Genus Protocirrineris Czerniavsky, 1881

Type Species: Cirratulus tenuisetis Grube,

1860, designated by Hartman (1959)

Diagnosis (after Blake, 1996; Blake &

Magalhães, 2017): Prostomium bluntly coni-

cal to wedge-shaped, with or without eyespots.

Body nearly round in cross section, with dis-

tinct segments. Grooved tentacular filaments

few, arising singly or in paired groups on sev-

eral anterior chaetigers following chaetiger 1;

individual tentacular filaments often arranged

in longitudinal rows. Branchiae occurring sin-

gly, usually first present from segments with

tentacular filaments, sometimes on more ante-

rior chaetigers. Chaetae all capillaries. Pygidi-

um a simple lobe.

Protocirrineris baiana sp. nov.

Fig. 1 A-D

Material examined: Northern Brazil,

Todos os Santos Bay, Paraguaçu estuary, Sta-

tion 1, 12°50’10.5” S, 38°48’04.8” W, May

2005, 16 m depth, two incomplete specimens

of 36 and 38 chaetigers; two complete speci-

mens, Holotype: a complete specimen, 23

mm long and 0.25 mm wide for 123 chaetigers

(MZUSP 3602); Paratypes: one complete

specimen, 19 mm long and 0.2 mm width ante-

riorly and 0.1 mm width posteriorly for 136

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

chaetigers (MZUSP 3603). Todos os Santos

Bay, Paraguaçu estuary, Sta. 1, 12°50’10.5” S,

38°48’04.8” W, December 2005, 16 m depth,

two incomplete specimens.

Additional material: several fragmented

specimens from the same locality and date for

Scanning Electronic Microscope.

Description: Holotype complete, 23 mm

long, 0.25 mm wide for 123 chaetigers. Body

in cross section more or less quadrangular

anteriorly, inflated dorsally and grooved ven-

trally. Anterior region laterally expanded with

shoulders. Dorsal crest from chaetiger 5 on

crowded body segments. Posterior region with

a ventral groove, dorsally and laterally expand-

ed; abruptly tapering to simple pygidium and

terminal anus (Fig. 1A, C). Color in alcohol

pale tan, with pink or orange specks. Tentacular

cirri and branchiae paler than body.

Prostomium very short, conical, tucked under

peristomium; eyespots absent; nuchal organs not

observed. Peristomium triannulated, with 3-4 sub

annulations in each annulus. First four chaetigers

as long as peristomium also with a sub-annula-

tion. Tentacular filaments from chaetigers 3-4 in

two groups of about 5-6 filaments each; branchiae

from chaetiger 1, right above notopodia and pres-

ent throughout (Fig. 1A, B).

Noto- and neuropodia well-separated ante-

riorly; in first 12-20 chaetigers, notopodium

more dorsal then becoming more lateral. Neu-

ropodia ventrally positioned throughout. Cap-

illary chaetae short, about 12-15 notopodial

capillaries and 8-10 neuropodial capillaries in

anterior segments. Posterior segments bearing

five notochaetae and two neurochaetae. Capil-

laries with a golden reflection, ending in flat

blade; one side of blade denticulate under SEM

(Fig. 1D).

Fig. 1. Protocirrineris baiana sp. nov. A: General view, anterior region. B: Anterior region; C: Posterior region and

pygidium; D: Capillary chaetae.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

Methyl Green Staining Pattern: bran-

chiae, tentacular filaments and ventrum stained

intense blue; prostomium and peristomium

stained pale blue.

Habitat: This species was collected in

shallow subtidal regions (16 m) from estua-

rine environments with variable salinities

(30-33) and sediments composed of predomi-

nantly coarse sand and granules (see Barros

et al., 2008).

Etymology: The name is referred to the

type locality, Baía de Todos os Santos (Todos

os Santos Bay), and ‘baiana’ refers to those

born in the Bahia state in which this Bay

is located.

Remarks: The achaetigerous region of

Protocirrineris baiana sp. nov. is very distinct

from other species of the genus by having a

conical anterior region, with a short prosto-

mium retracted into the peristomium. The

peristomium has three distinct annuli each with

3-4 sub annulation being readily distinct from

the peristomium of P. camamuensis sp. nov.,

which is shorter, triannulated but lacks distinc-

tive sub-annulations.

The four anterior chaetigers are as long

as the peristomium and they have second-

ary annulations. The first pair of branchiae

is anterior to the tentacular filaments arising

from chaetiger 1. Only two other species have

branchiae originating before the tentacular fila-

ments: P. socialis Blake, 1996 from California

and P. magalhaesi Elías et al., 2019 from South

Africa. The tentacular filaments of P. baiana

sp. nov. are not organized into rows but in two

groups of 5-6 filaments each. The posterior

region of the new species is inflated, only com-

parable with P. magalhaesi, but the anus is ter-

minal, while in the South African species it is

dorsal. Capillary chaetae are very short, stout,

denticulate in one side of the blade in P. baiana

sp. nov. For further comparison see Table 1.

Protocirrineris camamuensis sp. nov.

Fig. 2 A-D

Material examined: Northern Brazil,

Camamu Bay, Station 6S2A, 13°45’23.9” S,

39°04’50.6” W, 3.4 m depth, July 2005, Holo-

type: a single complete specimen (broken in

two parts during staining process), 20mm long

for 95 chaetigers (MZUSP 3604).

Description: Holotype complete, 20 mm

long for 95 chaetigers; quadrangular in cross

section, flattened dorso-ventrally with inflat-

ed dorsum and grooved ventrum. Chaetigers

crowded throughout. Parapodia forming shoul-

ders after tentacular filaments, reaching maxi-

mum width at chaetiger 35. Posterior region

rounded, laterally expanded, ending in a coni-

cal section of several crowded chaetigers, with

several sub-annulations; pygidium conical with

anus terminal. Color in alcohol tan, with whit-

ish branchiae and tentacular filaments.

Prostomium short and wide; barely pro-

truding from peristomium; eyespots absent and

nuchal organs not observed. Peristomium short,

subequal in length to four anterior chaetigers,

with three same-sized annuli. Tentacular fila-

ments short (possibly regenerating) rising from

chaetigers 3-4, organized into two groups of

5-6 filaments each. Branchiae from chaetiger

1, dorsal to notopodium, present to end of the

body but sporadically (Fig. 2A).

Noto- and neuropodia widely separated;

notopodia lateral and neuropodia ventral. Para-

podia forming distinct shoulders posterior to

tentacular filaments (Fig. 2B). Capillary chae-

tae present though out noto- and neuropodia.

Capillaries finely denticulated on one side of

blade (Fig. 2D). Notopodia with 10-20 capil-

laries per ramus, neuropodia with 12-15 capil-

laries per ramus in anterior region, decreasing

in number posteriorly. Parapodial tori slightly

projecting from body wall posteriorly and cap-

illaries shorter than anteriorly.

Methyl Green Staining Pattern: Entire

body stained with uncolored patches in the

ventrum of the peristomium, dorsal region and

parapodial shoulders, and the posterior cone

where the anus opens.

Remarks: Protocirrineris camamuensis

sp. nov. is clearly distinct from all other known

Protocirrineris species. The robust body is

very different from most Protocirrineris spe-

cies that usually have a slender body. The

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

general shape of the body, flattened and later-

ally expanded, is an unusual feature in Proto-

cirrineris. The only similarity is with one (out

of the two) P. socialis Blake, 1996 morphot-

ypes described by Blake (1996). However, the

pattern of tentacular filaments is different in

these two species. The presence of branchial

filaments from chaetiger 1 is shared with a few

species such as P. baiana sp. nov., P. socialis

from California and P. magalhaesi Elías et al.,

2019 from South Africa. A short prostomium is

present in P. baiana sp. nov., although in that

species is acute, and in P. camamuensis sp. nov.

is rounded and flattened. The posterior region

being laterally enlarged and expanded is unique

in the genus. The neuropodia are ventral from

the first chaetiger to the last similar to in P.

baiana sp. nov. See also Table 1.

Habitat: This species was collected with-

in Camamu Bay in shallow water estuarine

sediments. Sediment in the site is composed

of sand (79.9 %) and silt-clay (11.1 + 8.95 %,

respectively). The site has high values of Mn

and Fe and is assumed to have lowered the

diversity and abundance of the benthic assem-

blage (Hatje et al., 2008).

Etymology: This species is named after its

type locality Camamu Bay.

Protocirrineris sp. A

Fig. 3 A-D

Material examined: Northern Brazil,

Todos os Santos Bay, Paraguaçu estuary, Sta.

1, 12°50’10.5” S, 38°48’04.8” W, December

2005, 16 m depth, a single incomplete speci-

men, about 76 mm long for 85 chaetigers.

Additional material: several incomplete

fragments analyzed by SEM.

Fig. 2. Protocirrineris camamuensis sp. nov. A: anterior region; B: posterior region and pygidium; C: cross section in

thoracic region, showing long and abundant notochaetaes and the ventral neuropodium with short and scares chaetae.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

Description: Examined specimen incom-

plete; body quadrangular in cross-section

anteriorly, dorsally round and ventrally flat

with ventral groove. Color pale tan, with

iridescent spots, tentacular filaments and

branchiae whitish.

Prostomium short, triangular and flattened,

with two front-lateral dark spots (eyespots or

colored nuchal organs?). Peristomium tri-annu-

lated, as long as first five chaetigers. Tentacular

filaments in two groups (6-8 filaments each)

between chaetigers 3-4. Branchiae from chae-

tiger 1, dorsal to notochaetae and in posterior

part of segment.

Noto- and neuropodia separated; separa-

tion between noto- and neuropodia becom-

ing wider from chaetigers 6 to 21, forming

distinct shoulders. Neuropodia almost ventral

in anterior region. Capillary chaetae present

throughout, 15-20 capillaries in notopodia,

10-15 in neuropodia in anterior region, decreas-

ing progressively to end of body. All specimens

composed of anterior fragments, posterior end

and pygidium not known.

Methyl Green Staining Pattern: No clear

pattern of stain was observed.

Protocirrineris sp. B

Fig. 4 A-D

Material examined: Northern Brazil,

Todos os Santos Bay, Paraguaçu estuary, May

2015, Station 1 (3 incomplete specimens),

12°50’10.5” S, 38°48’04.8” W; December

2005, 16m depth; Station 5, 12°45’50.2” S,

38°52’35.5” W, three incomplete specimens;

Fig. 3. Protocirrineris sp. A. A: Schematic draws of anterior region and cross section. B: SEM of the anterior region. Note

the long and abundant capillaries in both noto- and neuropodium. C: Detail of capillary chaetae stout and finely striate in

one edge. D: Detail of capillary chaetaes finely denticulate with small spaced teeth and tip slightly raised, all around its

circumference.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

December 2005, Station 1, 12°50’10.5” S,

38°48’04.8” W, a single incomplete specimen

55 mm long for 63 chaetigers.

Additional material: several fragments

analyzed by SEM.

Description: All examined specimens’

incomplete posteriorly; longest anterior frag-

ment with 63 chaetigers. Body oval in cross

section, dorsally inflated, with a shallow ven-

tral groove anteriorly, more deeply grooved

posteriorly; body laterally expanded forming

shoulders in anterior region, reaching maxi-

mum width in chaetigers 26-33, subsequent

chaetigers are not as wide and more crowded.

Color in alcohol pale tan.

Prostomium short, conical; eyespots

absent; nuchal organs not observed. Peristo-

mium with two annuli best distinguished later-

ally; dorsal crest present along peristomium,

continuing to chaetiger 4 with a shallow central

groove. First three chaetigers longer than sub-

sequent ones, as long as peristomium (Fig. 4).

Tentacular filaments from chaetigers 2-4 right

after crest formed from peristomium; filaments

organized in two groups numbering 10-17

filaments in each group. First pair of branchiae

on first chaetiger, dorsal to notopodium; sub-

sequent branchiae on same position and bran-

chiae present anteriorly.

Noto- and neuropodia close together, most

anterior segments with notopodia almost dor-

sally placed and close to tentacular filaments,

then becoming lateral after chaetigers 4-5.

Neuropodia lateral throughout. Notopodia and

neuropodia with numerous very long capillar-

ies, longer and more abundant in notopodia,

22-25, and 13-18 in neuropodia anteriorly,

decreasing to the middle and posterior region.

Capillaries with dense tabs attached and tip

raised, some capillaries shorter with blunt

denticulations, very dense on one side, more

scattered on rest (Fig. 4).

Methyl Green Staining Pattern: Bran-

chiae, tentacular filaments and ventrum stained

dark blue; prostomium and peristomium stained

a paler blue.

Fig. 4. Protocirrineris sp. B. A: Schematic draw of anterior region. B: SEM of anterior region with long capillary chaetae.

C: Long capillaries denticulate, with numerous and tight teeth, the tip is raised. D: Capillaries with numerous and tight teeth.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

Remarks: Protocirrineris sp. B is a very

distinct member of the genus by the presence of

a dorsal peristomial grooved crest that extends

back to chaetiger 4. Capillary chaetae are

longer and numerous in comparison to other

known Protocirrineris species. The pattern of

ornamentation is also quite different from the

other congeners. Similarly to other Brazilian

species of Protocirrineris, the branchial fila-

ments are first present from chaetiger 1, ante-

rior to tentacular filaments.

Habitat: Specimens were collected at

middle and lower regions of Paraguaçu estu-

ary at Stations 1 and 5 with salinities varying

from 23-30 and sediments composed of mostly

sand and coarse sand (Sta. 1) and fine grained

sand (Sta. 5).

DISCUSSION

Two new species of Protocirrineris are

herein described and characterized by the pres-

ence of tentacular filaments in two packs

between chaetigers 3 and 4. The other two mor-

photypes are composed of incomplete speci-

mens but they seem to be adult individuals and

to be morphologically distinct from the two

new species. The posterior end of the body

was missing and there are no references about

the presence of spines in the last chaetigers,

however the general aspect of the body sug-

gests that they lack spines or any other type

of modified chaetae. This is the reason why

we identified these worms as belonging to

Protocirrineris. The identification and formal

description of these morphotypes must await

for new, complete material, to confirm the exis-

tence of additional new species.

The genus Protocirrineris is present in

the southwestern Atlantic (P. angellicolatio

Elías & Rivero, 2009), northeastern Pacific (P.

socialis Blake, 1996), northwestern Pacific (P.

mascaratus Magalhães & Bailey-Brock, 2013),

France (P. tenuisetis Grube, 1860), south-

western Pacific (P. nuchalis (Ehlers, 1907)),

Indic (P. indicus Day, 1973), Mediterranean

(P. chrysoderma (Claparède, 1868) and P.

purgamentorum Lezzi, Çinar, & Giangrande,

2016) and South African (P. strandloperarum

and P. magalhaesi, Elías et al., 2019). The two

species and the two morphotypes described

here constitute new additions to the known

Protocirrineris species from the coast of Bra-

zil (Table 1). The geographical distribution of

most species is restricted to the type locality

with the exception of P. chrysoderma, which

has been reported to be widespread in the

Atlantic, Mediterranean and Pacific. Petersen

(1991), however, argues that P. chrysoderma

may be a complex of species and a closer look

at these morphologically similar taxa using

SEM imaging could reveal that may be a much

more species-rich taxon (Lezzi et al., 2016;

Magalhães, 2015). The evidence suggests that

Protocirrineris species may be more locally

endemic than previously known.

The table provided by Magalhães & Bai-

ley-Brock (2013) was modified to include the

new species and novel morphotypes described

herein as well as the new species described in

Elías et al. (2019) from South Africa in this

same volume (Table 1). We propose a table,

with the characters herein described for each

new species, to aid in the identification of any

new species of Protocirrineris encountered.

The twelve described species of Protocir-

rineris (the Table does not include the two

morphotypes) may be distinguished by the

segmental origin and the extent and number of

feeding tentacles. Tentacular filaments could

be in longitudinal rows or in packs occupying

2-3 anterior chaetigers (see Table 1). The origin

of branchiae is usually from the posterior-most

tentacle-bearer segments but these are some-

times difficult to see without the aid of SEM,

but still are considered a diagnostic character.

Here we add two species and two morphotypes

with branchiae in anterior chaetigers, plus the

existing species (P. socialis). Methyl green

staining patterns have been described for four

out of the six species but only Protocirrineris

mascaratus has a distinctive staining pattern.

The description of P. tenuisetis is insufficient

to confirm its identity and according to Hart-

man (1959) this species should be considered

as indeterminate.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

The ultrastructure of the capillary fibrils

might be a relevant character but it has only

been shown for P. mascaratus and P. angelli-

colatio and they appear to be distinct in width,

shape, and number of fibrils along the capillary

chaetal blade (compare Fig. 4.3.2C, D herein to

Elías & Rivero, 2009: Fig. 2E, F). The useful-

ness of capillary fibrils to cirratulid taxonomy

has been discussed for the bitentaculate genus

Aphelochaeta (Doner & Blake 2009; but also

see Elías, Rivero, & Orensanz, 2016) but a uni-

fying terminology is lacking (see Fig. 4, Elías

et al. 2019, in this volume).

The phylogenetic relationships within the

Cirratulidae have never been investigated in

detail. The lack of reliable characters is prob-

lematic in the understanding of the phylogeny

of this group, as it has been shown that most

of the characters used in species level descrip-

tions of multitentaculate cirratulids, i.e. seg-

mental origin of feeding tentacles and acicular

spines vary ontogenetically (e.g. Magalhães

& Bailey-Brock, 2010). A related issue is the

broad generic diagnoses currently in use. Over

the years, the systematic accounts have been

modified to include a wider interpretation of

what the characteristics of a species included

in a particular genus are so that a place could

be found for those species that did not seem to

fit anywhere. These vague definitions result in

species having been placed in a genus more for

convenience than to reflect evolutionary rela-

tionships (Tewary, 2015).

Biogeographically, the region of study is

particularly interesting because two species

and an additional two Protocirrineris mor-

photypes were found in two very close sites

on the coast of Brazil. Recently, two species

of this genus were described for the coast of

South Africa (Elías et al. 2019, in this volume),

which was justified by the existence of three

biogeographical regions on that coast, although

the species of Brazil belong to the same

biographical region.

The understanding about the diversity of

cirratulid species in Brazil is in its first steps.

Additional sampling across different habitats,

examination of museum collections on top of

examining samples through molecular analysis

are needed in order to confirm the identifica-

tion of widespread and cryptic species.

Ethical statement: authors declare that

they all agree with this publication and made

significant contributions; that there is no con-

flict of interest of any kind; and that we fol-

lowed all pertinent ethical and legal procedures

and requirements. All financial sources are

fully and clearly stated in the acknowledge-

ments section. A signed document has been

filed in the journal archives.

See Table 1 in supplementary material

a07v67n5-MS1

AKNOWLEDGEMENTS

SEM images were obtained by Móni-

ca Oppedisano from the Laboratorio de

Microscopía Electrónica of Facultad de Cien-

cias Exactas y Naturales of the Universidad

Nacional de Mar del Plata. The drawings

were made by R.E. Figures composition by

M.A.S.B. Two anonymous reviewers improved

the manuscript.

RESUMEN

Dos nuevas especies de Protocirrineris (Polychaeta:

Cirratulidae) de Brasil

Introducción: Brasil tiene más de 8000 km de costa,

pero sus invertebrados marinos aún son poco conocidos.

Los poliquetos cirratulidos han sido un grupo especial-

mente descuidado debido a su desafiante taxonomía.

Objetivos: Identificar morfotipos de Cirratulidae multi-

tentaculados y encontrar nuevos caracteres diagnósticos.

Métodos: En muestreo reciente en dos de las bahías más

grandes de Brasil, Bahía de Todos los Santos y Bahía

de Camamu, se colectaron cuatro morfotipos del género

multitentaculado Protocirrineris. Resultados: debido a la

falta de quetas modificada, la taxonomía de Protocirrineris

es un desafío y solo dos de estos morfotipos se describen

aquí como nuevas especies. Estas dos nuevas especies

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 67(S5) Supplement: S81-S91, July 2019

se caracterizan por tener filamentos tentaculares en dos

grupos sobre los quetígeros 3-4 y las primeras branquias

presentes de quetígero 1. Se diferencian entre sí en relación

con la naturaleza de la región sin quetas, región posterior,

ultraestructura de las quetas capilares y el patrón de tinción

con verde de metilo. Dos especímenes incompletos tam-

bién se consideran Protocirrineris, y se dan breves descrip-

ciones. Conclusiones: Se cita por primera vez la presencia

del género Protocirrineris en Brasil. Cuatro morfotipos del

género han sido encontrados en dos grandes estuarios de

Brasil. Dos especies nuevas para la ciencia son descriptas.

Palabras clave: Taxonomía, Protocirrineris, especies nue-

vas, Bahía de Todos los Santos, Bahía de Camamu, Norte

del Brasil.

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