Received: 11 February 2022 Accepted: 22 May 2022

DOI: 10.1111/jfb.15117

REGULAR ARTICLE FISH

Redescription of Anchoviella cayennensis (Puyo, 1945)

(Clupeiformes: Engraulidae), with the synonymization
of Anchoviella sanfranciscana Barbosa, Gomes da Silva,
da Rocha Araújo & Carvalho, 2017 and remarks on Anchoviella
perfasciata (Poey, 1860)

Lorena S. Agostinho1,2 | Marina V. Loeb3 | Fabio Di Dario4

1
Programa de Po
s-Graduação em Ciências
Ambientais e Conservação (PPG-CiAC), Abstract
Universidade Federal do Rio de Janeiro (UFRJ), Anchoviella cayennensis (Puyo, 1945) is a relatively small-sized, coastal and estuarine
Macaé, RJ, Brazil
2

s-Graduação em Ecologia
western Atlantic species of anchovy distributed from Suriname to southeastern Brazil.
Programa de Po
(PPG-ECO), Departamento de Ecologia The species is morphologically similar to Anchoviella perfasciata (Poey, 1860) from the
(DECOL-CB), Universidade Federal do Rio
Central and western North Atlantic, and it has been suggested in the literature that both
Grande do Norte (UFRN), Natal, RN, Brazil
3
Museu de Zoologia da Universidade de São are actually synonyms. The recently described Anchoviella sanfranciscana (Barbosa et al.,
Paulo, São Paulo, SP, Brazil 2017) was reported as endemic to the estuary of the São Francisco River, northeastern
4
Instituto de Biodiversidade e Sustentabilidade
Brazil. Most characters proposed as diagnostic for A. sanfranciscana in the description
– NUPEM/UFRJ, Universidade Federal do Rio
de Janeiro, Macaé, RJ, Brazil are, however, similar to the ones reported for A. cayennensis and A. perfasciata. To
determine the complex taxonomic scenario involving the three species, 24 morphomet-
Correspondence
Fabio Di Dario, Instituto de Biodiversidade e ric and 13 meristic characters of 171 specimens tentatively identified as A. perfasciata
Sustentabilidade – NUPEM/UFRJ,
(including the holotype) and A. cayennensis from the Central and South Atlantic in addi-
Universidade Federal do Rio de Janeiro,
Av. São José do Barreto 764, CEP 27965-045, tion to the holotype and 19 paratypes of A. sanfranciscana (total: 191 specimens) were
Macaé, RJ, Brazil.
analysed. The PCAs of morphometric characters indicate the existence of two groups,
Email: didario@gmail.com
which are recognized as A. perfasciata and A. cayennensis, with A. sanfranciscana pro-
Funding information
posed as a junior synonym of the later. Further evidence from gill arch dentition also
CAPES; FAPERJ; PROTAX/CNPq
indicates that A. perfasciata and A. cayennensis are distinct valid species. A redescription
of A. cayennensis is presented, with a neotype proposed for the species. Confirmation
of the identity of specimens attributed to A. cayennensis indicates that its southern
limit of distribution is in the Rio de Janeiro State, southeastern Brazil. An updated taxo-
nomic key for the estuarine and coastal Atlantic species of Anchoviella is also presented.

KEYWORDS
anatomy, anchovies, conservation, Engraulinae, taxonomic key, western Atlantic

1 | I N T RO DU CT I O N Anchovies are relevant to artisanal and industrial fisheries on a global scale

(e.g., Birge et al., 2021), with recent catches of just a single species,
The Engraulidae includes 17 genera and c. 170 species of the typically Engraulis ringens (eastern Pacific of South America), reaching up to 12 mil-
small- to medium-sized coastal and schooling fishes commonly known as lion tonnes (FAO, 2020). They are also relevant to the maintenance of
anchovies (Hata et al., 2022; Loeb et al., 2018; Whitehead et al., 1988). coastal ecosystems because anchovies typically form large schools and

J Fish Biol. 2022;1–18. wileyonlinelibrary.com/journal/jfb © 2022 Fisheries Society of the British Isles. 1
2
FISH
compose a substantial portion of primary consumers among coastal fishes
(James, 1988). The family is currently divided into two subfamilies, the
Coiliinae (6 genera, Indo-Pacific distribution) and Engraulinae (11 genera,
worldwide distribution; Whitehead et al., 1988; Grande & Nelson, 1985;
Di Dario, 2009; Bloom & Lovejoy, 2012; Lavoué et al., 2014). The
Engraulinae is particularly diverse in the New World, where 9 genera and
c. 73 valid species are known to exist (Whitehead et al., 1988; Fricke
et al., 2022). Phylogenetic relationships among members of the
Engraulinae remain contentious, with most genera probably representing
non-monophyletic assemblages (e.g., Bloom & Lovejoy, 2012).
Anchoviella Fowler 1911 is a New World genus of Engraulinae that
includes 15 valid species exclusively found in coastal marine and fresh-
water environments (Barbosa et al., 2017; Bloom & Lovejoy, 2012;
Loeb, 2012, 2013; Loeb et al., 2018; Van der Sleen & Bloom, 2018). In
the last comparative taxonomic review of the Engraulidae, Whitehead
et al. (1988) defined Anchoviella by the following combination of char-
acters: a moderately slender and compressed body [depth c. 4.75–5
times in SL (standard length)], greatest height at dorsal fin origin; maxilla
moderate, posterior tip bluntly rounded and posterior to anterior mar-
gin of pupil, also failing to reach preoperculum by at least c. one-fourth
of pupil diameter; and long and numerous gill rakers, usually more than
15 on the lower branch of the first branchial arch.
Six coastal marine or estuarine species of Anchoviella are currently
recognized as valid in the western Atlantic (Loeb et al., 2018). Anchoviella
cayennensis (Puyo, 1945) was described based on two specimens col-
lected in the lower portion of the Cayenne River, French Guiana. The
two syntypes of the species were likely not preserved and could not be
located at the MNHN Fish Collection where they might have been
deposited (Fricke et al., 2022; Whitehead, 1973). Anchoviella cayennensis
was described based on a few characters, with comparative material
mostly restricted regionally. Whitehead (1973) redescribed the species
based on a single 56.2 mm SL specimen collected in the mouth of the
Coppename River, Suriname, with the remark that “Puyo's original
description of Stolephorus cayennensis (…) is poor but seems to refer to
the species described here” (Whitehead, 1973:161). Whitehead et al.
(1988) subsequently provided some additional comparative anatomical
remarks on the species. Published records of A. cayennensis in the liter-
ature are currently restricted to South America, between French Guiana
and Espírito Santo, Brazil (e.g., Menezes & Figueiredo, 2003; Whitehead
et al., 1988). Nonetheless, the identification of A. cayennensis among
congeners remains difficult given the problematic scenario of the diag-
nostic characters proposed for the species.
The morphologically similar Anchoviella perfasciata (Poey, 1860), also
from the western Atlantic, was described based on specimens collected
off Cuba. The geographic distribution of A. perfasciata is reported as
restricted to parts of the western Central and North Atlantic, between
North Carolina and Panama and possibly off Venezuela. Records in the
Gulf of Mexico require verification (e.g., Hildebrand, 1963; Hoese &
Moore, 1998; McEachran & Fechhelm, 1998; Whitehead et al., 1988).
When comparing the two species, Whitehead et al. (1988) stated that
“(…) Anchoviella perfasciata and (…) A. cayennensis differ from all other
members of the genus in having the anal fin origin behind the dorsal fin
basin. Possibly cayennensis is merely a southern subspecies [of the
AGOSTINHO ET AL.
former]” (Whitehead et al., 1988:338). Actually, the origin of the anal fin
is also distinctly placed posteriorly in relation to the dorsal-fin base in
some South American freshwater species of the genus, such as Ancho-
viella carrikeri Fowler, 1941 (Amazon basin) and Anchoviella guianensis
(Eigenmann, 1912) (Orinoco and Amazon basins, also coastal between
Venezuela and Pará, northern Brazil), but the condition was indeed
restricted to A. cayennensis and A. perfasciata among coastal/estuarine
species of Anchoviella until the description of Anchoviella sanfranciscana
2017 (Barbosa, Gomes da Silva, da Rocha Araújo & Carvalho, 2017).
The complex taxonomic scenario involving A. perfasciata and
A. cayennensis was further complicated with the description of
A. sanfranciscana. The three species can hardly be differentiated in terms
of the external anatomical characters proposed as diagnostic in the litera-
ture, except for a slightly smaller number of rakers in the lower portion
of the first branchial arch in A. perfasciata (24–30 vs. 28–35 in
A. cayennensis and A. sanfranciscana; Whitehead et al., 1988; Barbosa
et al., 2017). Other characters proposed by Barbosa et al. (2017) for
A. sanfranciscana are rather problematic in terms of description. Likely as
a result of this situation, records of A. sanfranciscana are still restricted to
the 20 specimens that compose the type series collected in the estuary
of the São Francisco River, northeastern Brazil, to which the species was
regarded as endemic at the description.
Here, A. cayennensis is redescribed based on characters of exter-
nal and internal anatomy, and the validity of A. sanfranciscana and the
taxonomic status of A. perfasciata are discussed An identification key
for marine and estuarine species of Anchoviella of the Atlantic is also
provided based on literature and examined specimens.
2 | M A T E R I A L S A N D M ET H O D S
2.1 | Specimens
A total of 191 specimens of Anchoviella from Brazil, Cuba, French Gui-
ana, Martinique and the United States were analysed. Specimens were
tentatively identified based on literature (Whitehead, 1985; Whitehead
et al., 1988; Loeb, 2012, Loeb, 2013; Loeb & Figueiredo, 2014; Barbosa
et al., 2017; Loeb et al., 2018) and geographic distribution (Figure 1),
resulting in the pre-identification of 131 specimens as A. cayennensis
(South America) and 40 as A. perfasciata (Central and North America),
including its holotype (MCZ 17965). The type series of A. sanfranciscana,
which includes the holotype and 19 paratypes, all collected in the estu-
ary of the São Francisco River, Brazil (Barbosa et al., 2017), was also
examined. As mentioned in the Introduction, the two syntypes of
A. cayennensis are lost. The specimen examined by Whitehead (1973;
RMNH 26294) was not available for this study; nonetheless, a topotype
of A. cayennensis (MNHN-IC 2005-2320) was included in the analyses.
2.2 | Morphological data
Counts and measurements were taken according to Hubbs and Lagler
(1947) and Loeb et al. (2018). SL and head length (HL) measurements
AGOSTINHO ET AL.
FISH 3

F I G U R E 1 Collection
localities of 191 specimens
examined in this study,
tentatively identified as
Anchoviella perfasciata (blue
circles), Anchoviella cayennensis
(yellow circles) and the type
series of Anchoviella
sanfranciscana (red circle)

are expressed in millimetres. Measurements of the body are
expressed as a percentage of SL, except subunits of the head that
are expressed as a percentage of HL (Table 1). Meristic data are pro-
vided in Table 2 and in the redescription of A. cayennensis, with an
asterisk indicating counts of A. cayennensis MNHN-IC 2005-2320,
proposed here as the neotype for the species. Frequency (number of
specimens) of each count is given in parentheses in the “Diagnosis”
and “Description” sections.
Five specimens identified as A. cayennensis (NPM1883, 2;
NPM1884, 1; NPM3265, 2; 102.2–141.2 mm SL), four identified as
A. perfasciata (UF 220202, 65–78 mm SL) and one paratype of
A. sanfranciscana (INPA-ICT 053280) were cleared and stained for visual-
ization of bones and cartilages (Dingerkus & Uhler, 1977; Song &
Parenti, 1995; Taylor & Van Dyke, 1985). Osteological observations were
complemented by X-ray images obtained using a Digital Faxitron X-Ray
Machine model LX-60 of 13 specimens identified as A. cayennensis and
10 as A. perfasciata, in addition to 10 specimens of the type series of A.
sanfranciscana, including the holotype (INPA-ICT 053278).
Osteological and lateral line terminology follow Nelson (1967,
1970), Grande (1985), Grande and Nelson (1985), Di Dario (2002,
2004) and Carvalho et al. (2013).
2.3 | Morphological analyses
PCAs were used to investigate the morphometric patterns of the species
studied. Two PCAs were performed with the morphometric data. The
first PCA included a non-standardized matrix of 191 specimens of
4
FISH AGOSTINHO ET AL.

T A B L E 1 Morphometric data of 191 specimens identified as Anchoviella cayennensis (n = 131); Anchoviella perfasciata (n = 40), including the
holotype (MCZ 17965); and the type series of Anchoviella sanfranciscana (n = 20), including the holotype and 19 paratypes (S.D.)

A. cayennensis A. perfasciata A. sanfranciscana

Other specimens Other specimens Paratypes

MNHN-IC
2005–2320 Range Mean S.D. Holotype Range Mean S.D. Holotype Range Mean S.D.

SL (mm) 98.5 54–145.5 112.4 – 79.4 42.8–81 63.1 – 105 97–114.7 106.8 –
HL (mm) 22.3 13.7–32.1 25.4 – 19.4 10.4–21.1 15.8 – 23.5 22.2–26 18.6 –
% of SL
Body depth 15.8 11.3–19.8 17.1 1.6 13.8 12.7–17.4 14.5 1.2 19 16.5–19.6 18.6 0.7
Caudal peduncle 7.6 6.5–10.1 7.8 0.4 8 6.5–8.6 7.6 0.4 7.9 6.9–9.7 8.5 0.6
depth
Head length 22.6 19.8–25 22.7 0.8 24.5 23.6–26.3 25.1 0.6 22.3 20.9–24.3 22.7 0.7
Dorsal-fin base 9.7 8.6–15.1 11.8 1 10.2 9.8–14.4 12.3 0.9 12.2 9.8–14.4 12.1 0.9
length
Anal-fin base length 11.2 9.3–14.3 11.5 1 13.1 12.5–18 14.6 1.2 12.9 10.4–12.9 11.3 0.8
Pectoral-fin length 15.4 12.7–19.7 14.9 1 11.2 12.3–15.4 13.4 0.8 14.8 13.5–16.4 15.1 0.7
Pelvic-fin length 8.6 7.4–12.4 9.3 1.2 6.6 6.72–10.9 8.1 0.8 10 8.7–11.5 9.8 0.6
Pre-dorsal length 49.8 45.6–56.3 52.5 1.9 51.6 47–56.1 52 1.9 50.1 45.3–55 52.3 2
Pre-pectoral length 23.1 21.4–30.3 23.8 1.1 23.7 24.2–28.2 26.4 0.8 24.1 19.1–25.5 24 1.4
Pre-pelvic length 43.1 40.6–53 43.1 3.2 46.1 40.3–47.5 44.6 1.8 44.2 39–45.1 43.7 2.2
Pre-anal length 65.4 58.7–75.6 67.7 2.4 64 61–67 64.3 1.7 67.9 59.4–70.2 67.7 2.3
Pectoral-fin axillary 15.4 9.4–17.4 13.5 5.6 10.1 7.1–12.4 10.3 1.2 – – – –
scale length
Pelvic-fin axillary 8.9 7.7–11.1 9.3 0.9 – – – – – – – –
scale length
Distance between 4.7 2.8–10 6.4 1.7 10.1 4–13.8 7.5 2 8.1 5.7–10.1 7.5 1.1
pectoral-fin tip
and pelvic-fin
base
Distance between 12.8 8.5–20 14.7 2.2 11.5 8.6–14 11 1.5 14.2 10.7–18.1 14.8 1.6
pelvic-fin tip and
anal-fin base
% of HL
Snout length 14.6 13–25 18.6 2.6 14.4 15.7–24.1 18.6 2.1 17.8 17.4–21.2 18.9 1
Upper-jaw length 56.5 56.1–69.4 65.1 1.9 69.9 69.2–77.3 74.1 1.9 67.6 62.5–67.6 65 1.2
Upper-jaw depth 4.9 3.8–8.7 5.2 0.6 4.3 6.1–8.3 7.3 0.8 5.5 4.6–6.3 5.6 0.3
Upper-jaw length 19.1 13.2–25.4 17 1.9 19.4 17.6–27 23.5 2.1 19.5 15.3–21.9 18.4 1.8
beyond posterior
margin of orbit
Orbit diameter 29.4 28–37.5 32.4 1.9 31.7 30.6–37.8 33.6 2.3 29.7 27.7–34.5 31.7 1.5
Post-orbital distance 50 44–58.6 52.2 2.5 48.4 46.7–56 50.4 2.1 52.7 46.6–56.6 52.5 2.1
Interorbital width 25.3 19.8–31.9 24.2 1.6 23.5 16.5–25.1 21.5 1.4 25.5 22.9–26.2 24.8 0.9

Note: HL, head length; SL, standard length.

Anchoviella directly examined in this study as rows and all 22 morphomet-
ric data as columns (Table 1). The second PCA was performed with data
from those same 191 specimens in addition to data from A. cayennensis
RMNH 26294 examined by Whitehead (1973) in the redescription of
the species. As this specimen was not examined in the current study,
only the following 10 morphometric data reported by Whitehead (1973)
were included in the second PCA: body depth, anal-fin base length,
pectoral-fin length, pelvic-fin length, pre-pelvic distance, pre-anal dis-
tance, HL, snout length, upper-jaw length and orbit diameter. All analyses
were performed using the R Program (www.r-project.org) with assistance
of the packages “Factominer,” “Factoextra” and “Ggplot2.” (Everitt &
Hothorn, 2011; Venables et al., 2019).
AGOSTINHO ET AL.
2.4 | Institutional abbreviations

gica da
Institutional abbreviation are as follows: CIUFES, Coleção Ictiolo
Universidade Federal do Espírito Santo, Espírito Santo, Brazil; INPA,
Instituto Nacional de Pesquisas da Amazônia, Amazonas, Brazil; MCZ,
Museum of Comparative Zoology, Harvard University, Massachusetts,
USA; MNHN, Muséum National d'Histoire Naturelle, Paris, France; MNRJ,
Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro,
Brazil; MPEG, Museu Paraense Emilio Goeldi, Pará, Brazil; MZUSP, Museu
de Zoologia da Universidade de São Paulo, São Paulo, Brazil; MZFS,
Divisão de Peixes, Museu de Zoologia da Universidade Estadual de Feira
de Santana, Feira de Santana, Brazil; NPM, Coleção de Peixes do Instituto
de Biodiversidade e Sustentabilidade, Universidade Federal do Rio de
Janeiro, Rio de Janeiro, Brazil; RMNH, Naturalis Biodiversity Center, Lei-
den, Netherlands; UF, Florida Museum of Natural History, Florida, USA;
UFS, Universidade Federal de Sergipe, Aracaju, Brazil; and USNM, National
Museum of Natural History, Smithsonian Institution, Washington, D.C.
2.5 | Ethical statement
This study used only ethanol-preserved specimens deposited in
museums and did not involve animal experimentation.
3 | RESULTS
A. cayennensis (Puyo, 1945): Figures 2 and 3; Tables 1 and 2.
Stolephorus cayennensis Puyo, 1945: 101–103, figure 1-I [original
type locality: “rivière Cayenne à hauteur de la pointe Macouria, à envi-
ron quatre kilomètres de son embouchure” (=Cayenne River in
Macouria, about 4 km from its mouth), French Guiana; also reported
from “rivière Mahury et le fleuve Kourou” (=Mahury and Kourou Riv-
ers)]; Puyo, 1949: 157, figure 80 (list of fishes from French Guiana;
description republished, no new observations).
Anchoviella cayennensis (Puyo, 1945): Hildebrand, 1963: 212 [new
combination; redescription based on published accounts and figures by
Puyo (no specimens examined) and taxonomic key]; Whitehead, 1973:
161, figures 61 and 62 (redescription based on one specimen from Suri-
name); Whitehead & Bauchot, 1986: 49 (information on the type-series
specimens); Whitehead et al., 1988: 330 (diagnosis, biology and distribu-
tion; possible subjective junior synonym of A. perfasciata); Camargo &
Isaac, 2001: 140 (list of estuarine fishes of northern Brazil); Kullander &
Ferraris Jr. 2003: 40 (list of freshwater fishes of Central and South Amer-
ica); Nizinski & Munroe, 2003: 766 (list of fishes from western Central
Atlantic); Barbosa et al., 2017: 167 (list of comparative material);
Menezes & Figueiredo, 2003: 39 [as Anchoviella cayenensis (wrong spell-
ing), list of marine fishes of Brazil]; Loeb, 2012: 18 [as Anchoviella
cayenensis (wrong spelling), list of comparative material and taxonomic
key]; Loeb & Figueiredo, 2014: 35 [as Anchoviella cayenensis (wrong spell-
ing), list of comparative material]; Loeb et al., 2018: 2 [as Anchoviella
cayenensis (wrong spelling), list of valid species of Anchoviella]; Birge et al.,
2021 (Supplementary Data, conservation status).
FISH 5
Anchoviella victoriae Hildebrand & de Carvalho, 1948: 292 (type

ria, State of Espírito Santo, Brazil); Carvalho, 1951: 51, fig-
locality: Vito
ure 1 (list of Brazilian species of Anchoviella); Santos, 1952: 75 (list of
marine fishes of Brazil); Whitehead, 1973: 161 (junior synonym of
A. cayennensis); Menezes, 1974: 216 (holotype information); White-
head et al., 1988: 330 (junior synonym of A. cayennensis).
Anchoviella sanfranciscana Barbosa et al., 2017: 162, figures 2, 3
and 5 (type locality: northeastern Brazil, São Francisco River estuary
between the cities of Brejo Grande, state of Sergipe, and Piaçabuçu,
state of Alagoas, Brazil); Vicente et al., 2020: 3 (mentioned as a possi-
ble junior synonym of A. cayennensis based on unpublished data);
Birge et al., 2021 (Supplementary Data, conservation status).
3.1 | Neotype
As indicated in the Introduction, syntypes of Stoleporus cayennensis
Puyo, 1945, either have been lost for decades or have never been
deposited in the MNHN Fish Collection, as first noticed by Whitehead
(1973). This conclusion is further supported by a recent visit to the
MNHN Fish Collection by one of the authors (M.V.L.), who searched
for the specimens without success. Given the complex taxonomic situa-
tion involving A. cayennensis and other similar species of Anchoviella, a
neotype is designated to define the nominal taxon more objectively
(ICZN, 1999): MNHN-IC-2005-2320, 98.5 mm SL, Atlantic Ocean,
Caribbean Sea, Vieux Port, Cayenne, French Guiana, 4
550 1.200 N,
52
70 58.800 W (new type locality), 19 March 2002, M. Leopold.
3.2 | Non-type specimens examined

bay,
Brazil, Pará: MPEG 6304 (1, 115.0 mm SL), Jubim beach, Marajo
Amazon River, 45
480 6300 S, 48
300 4100 W, 26 May 1982; MPEG

bay, Amazon River,
6306 (1, 115.7 mm SL), Jubim beach, Marajo
45
480 6300 S, 48
300 4100 W, 26 June 1982. Alagoas: MZUSP
113730 (2, 115.2–117.0 mm SL), Penedo, São Francisco River, 20 Jan-
uary 2013. Sergipe: UFS 1132 (19, 92.3–122.0 mm SL), Cabeço town,
São Francisco River, 10
300 2.600 S, 36
240 2800 W, September 1975;
UFS 1131 (1, 114.5 mm SL), continental shelf off Sergipe, 2000; UFS
1128 (2, 97.5–115.1 mm SL), continental shelf off Sergipe,
10
520 3300 S, 36
560 3200 W, August 1999; UFS 1129 (2, 96.2–
115.9 mm SL), continental shelf off Sergipe, 10
500 5300 S,
36
540 5300 W, November 1988; UFS 1130 (3, 58.2–125.1 mm SL),
continental shelf off Sergipe, 2000; CIUFES 3001 (3, 102.7–

polis, São Francisco River, 10
180 56.400 S,
105.7 mm SL), Neo
36
340 28.200 W, 14 September 2013; MZUSP 11579 (1, 86.3 mm
SL), Aracaju, August 1961. Bahia: MNRJ 8204 (1, 116.0 mm SL), Caixa
Prego, 12 December 1944; MZUSP 11583 (1, 113.8 mm SL), Pontal
beach, Ilhéus, 25 October 1971; MZUSP 11582 (8, 92.0–118.3 mm
SL), Malhadinho beach, Ilhéus, 25 October 1971; MZUSP 11580
(1, 107.0 mm SL), Malhadinho beach, Ilhéus, 26 October 1971; MZFS
12712 (4, 117.0–118.8 mm SL), in front of “Ponta da Tulha,”
Ilhéus; MZFS 9860 (2, 90.2–92.4 mm SL), Malhado beach, Ilhéus.
6
FISH AGOSTINHO ET AL.

T A B L E 2 Meristic data of 191 specimens identified as Anchoviella cayennensis (maximum n = 131); Anchoviella perfasciata (maximum n = 40),
including the holotype (MCZ 17965); and the type series of Anchoviella sanfranciscana (maximum n = 20), including the holotype and 19
paratypes

Dorsal-fin rays

ii iii 9 10 11 12 13 14 N

A. cayennensis 4 127* 1 3 18 62* 41 4 131

A. perfasciata 2* 33 4* 30 6 35
A. sanfranciscana 20* 4 15* 1 20

Pectoral-fin rays

i ii 10 11 12 13 14 15 N

A. cayennensis 126* 1 2 23 36 57* 4 126

A. perfasciata 37* 3 2 5 31* 2 37
A. sanfranciscana 20* 10* 9 20

Pelvic-fin rays

i ii 5 6 7 N

A. cayennensis 127* 1 3 110* 14 127

A. perfasciata 40* 1 37* 1 40
A. sanfranciscana 20* 20* 20
Anal-fin rays

ii iii 11 12 13 14 15 16 N

A. cayennensis 2 126* 9 51 61 7* 128

A. perfasciata 5* 35 16* 5 13 40
A. sanfranciscana 20* 2 8 9* 20

Caudal-fin rays: upper portion

iiii 9 10 11 N

A. cayennensis 130 61 56 13 130

A. perfasciata 39 18 15 6 39
A. sanfranciscana 20* 10* 9 1 20

Caudal-fin rays: lower portion

iiii 9 10 11 N

A. cayennensis 130 55 60 15 130

A. perfasciata 39 16 7 6 39
A. sanfranciscana 20* 10* 7 3 20

Total gill rakers: first branchial arch

45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 N

A. cayennensis 2 3* 6 9 20 20 23* 14 10 12 7 1 1 128

A. perfasciata 2 6 9 8 8* 3 3 39
A. sanfranciscana 1 1 3 4 3 2 3 2 1 20
Upper-portion gill rakers: first branchial arch
19 20 21 22 23 24 25 26 27 28 N
A. cayennensis 1 11* 17 19 24 31 18 6 2 2 128
A. perfasciata 2 13 14* 10 1 39
A. sanfranciscana 1 4 4 5* 5 1 20

Lower-portion gill rakers: first branchial arch

25 26 27 28 29 30 31 32 33 34 35 N

A. cayennensis 2 3 27* 36 24 30 7 2 128

AGOSTINHO ET AL.
FISH 7

TABLE 2 (Continued)

Lower-portion gill rakers: first branchial arch

25 26 27 28 29 30 31 32 33 34 35 N

A. perfasciata 6 12 9 9* 3 39
A. sanfranciscana 5 6 2 6* 1 20

Precaudal vertebrae Caudal vertebrae Total vertebrae

23 24 17 18 19 20 41 42 43 N

A. cayennensis 2 11* 9* 2 2 8* 2 4 13
A. perfasciata 7* 3 1 7* 2 7 3* 10
A. sanfranciscana 2 8* 6 3* 1 7 3* 10

Pre-dorsal bones

10 N

A. cayennensis 13* 13
A. perfasciata 10* 10
A. sanfranciscana 10* 10

Note: “*” indicates MNHN-IC 2005-2320 and holotypes of A. perfasciata and A. sanfranciscana, and “i” indicates number of unbranched fin rays.

F I G U R E 2 (a) Anchoviella perfasciata (Poey, 1890), holotype, MCZ

17965, 79.4 mm SL (Museum of Comparative Zoology, Harvard
University); (b) Anchoviella cayennensis (Puyo, 1945), neotype, MNHN-
IC-2005-2320, 98.5 mm SL; (c) Anchoviella sanfranciscana, holotype,
F I G U R E 3 X-ray images of (a) Anchoviella perfasciata, holotype,
INPA-ICT 053278, 105.0 mm SL (after Barbosa et al., 2017)
MCZ 17965, 79.4 mm SL; (b) A. perfasciata, UF 208245, 64.0 mm SL;
(c) Anchoviella cayennensis (Puyo, 1945), neotype, MNHN-IC-
2005-2320, 98.5 mm SL (MNHN, ICHTYOLOGIE-J. PFLIGER and
Espírito Santo: MZUSP 108171 (3, 133.0–135.0 mm SL), Vila Z. GABSI); (d) A. cayennensis, CIUFES 1131, 110.0 mm SL;
(e) Anchoviella sanfranciscana, holotype INPA-ICT 053278,
Regência, Doce River, 15 September 2005; MZUSP 11581 (4, 84.4–
105.0 mm SL
89.6 mm SL), Vitoria; CIUFES 130808 (22, 93.6–121.0 mm SL),
Itapina, Doce River, 19
310 58.900 S, 40
370 59.8 00 W, 9 May 1980;
CIUFES uncatalogued (1, 127.5 mm SL), Regência, Doce River,
19
380 02.100 S, 39
490 10.8 00
W, July 2016; MZUSP 60368 14774 (1, 135.3 mm SL), São Fidélis, Paraíba do Sul River, 21
400 S,
(23, 104.0–145.5 mm SL), Colatina, Doce River, 1 May 1944. Rio de 41
450 W, November 1989; MZUSP 109205 (1, 126.9 mm SL), São

i, 6 October
Janeiro: MNRJ 11126 (1, 54.0 mm SL), Itaipu beach, Nitero Fidélis, Paraíba do Sul River, 21
380 0000 S, 041
450 3000 W, 29 March
1978; MNRJ 37790 (2, 108.7–126.7 mm SL), Capixete Island, 2010; NPM 1883 (6, 91–141.6 mm SL), São Fidélis, Paraíba do Sul
Itaocara, Paraíba do Sul River, 29 October 2010; MNRJ 17869 River, 21
370 60.000 S, 41
380 16.000 W, 29 April 2012; NPM 1884
(3, 123.5–145.0 mm SL) São Fidélis, Paraíba do Sul River, 1996; MNRJ (5, 101.1–113.1 mm SL), Campos dos Goytacazes, Paraíba do Sul River,
8
FISH
F I G U R E 4 Lower portion of the branchial arches of (a) Anchoviella
cayennensis (NPM 1883) and (b) Anchoviella perfasciata (UF 220202) in
oral view, and upper portion of the branchial arch of A. cayennensis
(NPM 1883) in (c) oral and (d) dorsal views. Abbreviations: ac4:
accessory element of ceratobranchial 4; bh: basihyal; b1-4:
basibranchials 1–4; c1-5: ceratobranchials 1–4; e1-4: epibranchials 1–
4; go: gongyloid cartilage; h1-3: hypobranchials 1–3; i1-4:
infrapharyngobranchials 1–4. Branchial arch dentition is indicated in
light red. Gill rakers and filaments are not represented. Scale
bars = 1 mm
30 March 2012; NPM 3265 (7, 125.0–137.6 mm SL), São Fidélis, Par-
aíba do Sul River, 21
370 60.000 S, 41
380 16.000 W, 5 August 2013.
3.3 | Type series of A. sanfranciscana, junior
synonym of A. cayennensis
Holotype: INPA-ICT 053278 (106.0 mm SL), São Francisco River estu-
ary between cities of Brejo Grande, State of Sergipe, and Piaçabuçu,
State of Alagoas, northeastern Brazil, 10
240 44.100

0 00
36 27 28.4 W, A.G.S. Silva & A.R.R. Araújo, 12 April 2015. Para-
types: INPA-ICT 053279 (4, 105.5–114.7 mm SL), São Francisco River
estuary between the cities of Brejo Grande, State of Sergipe, and
Piaçabuçu, State of Alagoas, 10
240 44.100 S, 36
270 28.400 W, stated
as 12 April 2016 but likely collected with the holotype (12 April
2015). Sergipe: INPA-ICT 053280 (15, 103.6–128.0 mm SL), São
Francisco River estuary near Saramen village in the municipality of
AGOSTINHO ET AL.
F I G U R E 5 Head (after Whitehead, 1973) and inset of the otic
region of Anchoviella cayennensis (NPM 3265) showing the
openings of the recessus lateralis in left lateral view.
Abbreviations: acr?: possible accessory temporal sensory canal
opening to the chamber of the recessus lateralis; aor: anterior
opening to the chamber of the recessus lateralis; epo: epioccipital
at the floor of the pre-epiotic fossa; lfr: lateral wing of the frontal;
mor: middle opening to the chamber of the recessus lateralis; pa:
parietal; pfh: fossa on the pterotic that articulates with the
posterior condyle of the hyomandibula; por: posterior opening to
the chamber of the recessus lateralis; pto: pterotic; pfh: fossa on
the sphenotic that articulates with the anterior condyle of the
hyomandibula; sp: sphenotic; tf: temporal foramen; infraorbital
bones, preopercle and hyomandibula are not depicted in the inset.
Scale bar = 1 mm
Brejo Grande, 10
280 22.200 S, 36
250 02.600 W. A.G.S. Silva &
A.R.R. Araújo, 19 November 2015.
3.4 | Diagnosis
Anchoviella cayennensis is distinguished from congeners, except
A. juruasanga, A. guianensis, A. carrikeri, A. vaillanti and A. perfasciata
by the anal-fin origin just below or behind vertical through posterior
margin of last dorsal-fin ray base (vs. anal-fin origin anterior to vertical
through posterior margin of last dorsal-fin ray). Anchoviella cayennensis
can be distinguished from A. juruasanga, A. guianensis, A. carrikeri and
A. vaillanti by 28–35 gill rakers in the lower portion of the first bran-
chial arch (vs. 12–27). Anchoviella cayennensis is distinguished from
A. perfasciata by a reduced basibranchial dentition, consisting of a nar-
row median tooth plate over basibranchials 1–3, not expanded poste-
S, riorly over hypobranchials 2 (vs. a developed basibranchial dentition,
tooth plate over basibranchials 1–3 wide and expanded posteriorly
over hypobranchials 2; Figure 4a,b), and by the presence of a series of
delicate teeth over the lower portion of gill arches 1 and 2 (vs. teeth
absent; Figure 4a,b). Anchoviella cayennensis can also be distinguished
from A. perfasciata by 11–14 branched anal-fin rays (vs. 14–16) and
by a slightly smaller upper jaw, 56.1%–69.4% (mean 65.1%) HL vs.
69.2%–77.3% (mean 74.1%) HL.
AGOSTINHO ET AL.
3.5 | Description
Morphometric and meristic data of neotype and examined specimens
are summarized in Tables 1 and 2, including the type series of
A. sanfranciscana. Body elongated and slightly compressed laterally,
maximum height at vertical through dorsal-fin origin, c. five to eight
times in SL; a longitudinal silvery stripe along lateral of the body, from
mid- to upper-third of branchial opening to middle of caudal-fin base,
height of stripe at origin of dorsal fin 1–1.5 times eye diameter; upper
and ventral margins of lateral stripe with black pigment, more evident
at upper margin in preserved specimens; pre-dorsal bones 10(23),
total vertebrae including compound centrum (first preural + first ural
centrum + uroneural) 41(15)*, 42(5) or 43(3); precaudal vertebrae
23(4) or 24(19)*, caudal vertebrae 17(15)*, 18(5) or 20(3); maximum
body size 145.5 mm SL.
Head longer than deep, anterior tip gently pointed dorsally; snout
short, about half orbit diameter, two slit-like parallel nostrils on top of
snout at each side of head; mouth slightly inclined posteriorly, subter-
minal; upper jaw slightly curved, relatively short, 56.1%–69.4% (mean
65.1%) HL; posterior margin of upper jaw rounded, slightly projecting
beyond tip of second supra-maxilla, failing to reach anterior margin of
preopercle by half the pupil diameter; upper-jaw length beyond poste-
rior margin of orbit 13.2%–25.4% (mean 17.0%) of HL; a single row of
delicate, sharp, tapered curved teeth in the premaxilla, maxilla and
dentary; teeth in the maxilla slightly inclined backwards in the anterior
third of the bone, slightly inclined forward in the remaining structure;
a “gap” composed of a toothless area at the anterior portion of the
dentary; posterior projections of vomer with one to three minute
teeth; conical teeth present at the ventral border of the palatine, con-
tinuous to a similar row of teeth at the ventral border of the
ectopterygoid; small, conical teeth also forming a medial row on the
anterior half of the endopterygoid; first supramaxilla long, about
two-thirds of maxilla length and partially covered laterally by
infraorbital series; second supramaxilla also developed but smaller, a
little less than half maxilla length, posterior half scale-like, anterior
portion in the form of a spike; eyes large, orbit diameter 27.7%–
37.5% (mean 32.3%) of HL, dorsally located in relation to the longitu-
dinal line passing through the dorsal point of insertion of pectoral fin,
also visible in ventral and dorsal views; operculum length about twice
in height, lower margin oriented at an angle of c. 60
; posterior and
lower margins of preopercle forming an angle of c. 90
; exposed por-
tion of subopercle long and narrow; posterior border of gill opening
evenly rounded; pseudobranch present on inner side of operculum,
smaller than orbit diameter, 14.8%–23.3% HL; lateral line channels of
the panamensis-type (sensu Whitehead et al., 1988) on the operculum,
i.e., without a secondary preopercular branch projecting into the
opercle.
Recessus lateralis chamber developed, in the approximate shape
of a bell, tapered dorsally to the supraorbital canal, mostly covered lat-
erally by the lateral wing of the frontal; anterior opening of the
recessus, connected to infraorbital canal, mostly downward directed,
developed, located between the anterior (sphenotic) and posterior
(pterotic) sockets for articulation of the hyomandibula head (Figure 5);
FISH 9
lateral wall of the anterior opening formed by the lateral wing of the
frontal, medial and anterior walls delimited by the sphenotic, posterior
wall formed by a small flange of the pterotic that separates it from the
middle opening of the recessus; middle opening of the recessus, con-
nected to the preopercular canal, located immediately posterior to the
anterior opening, lateral wall formed by the lateral wing of the frontal
but also medially and anteriorly delimited by the pterotic; middle
opening of the recessus continuous with a scythe-like posterior open-
ing of the recessus at the posteroventral border of the lateral wing of
the frontal (possibly an expanded accessory canal opening to the
recessus chamber); posterior opening of the recessus developed,
oblong shaped, its height about four times its maximum length at
the middle of the opening; posterior opening connected dorsally to
the single branch shared anteriorly by the extrascapular and post-
temporal canals at the extrascapular bone, but also apparently con-
nected to superficial canals at the opercular region ventrally; posterior
opening of the recessus restricted by the pterotic, totally encircled by
a flange of the bone; pterotic at the posterior portion of the opening
forming an approximately triangular process with a posterodorsally
directed tip.
Prootic and pterotic bulla inconspicuous but fairly developed,
pterotic bulla laterally round, not bulging into pre-epiotic fossa, and at
the level of the recessus chamber floor in lateral view; temporal fora-
men exposed, not covered laterally by the frontal, shaped in the form
of an upside-down triangle; pre-epiotic fossa also developed, some-
what round in shape but bulged posteroventrally, mostly limited
anterodorsally by the parietal, anteroventrally by the pterotic, and
posteriorly by the epioccipital; a conspicuous circular opening at the
posteroventral region of the pre-epiotic fossa in the epioccipital, lead-
ing to the back of the skull; extrascapular canals at parietal not totally
encircled by bone, ossified segment of canals restricted to distal third
of the dorsal portion of the parietals; extrascapular canals at parietals
immediately posterior to posterior margin of frontals at the top of
cranium.
Dorsal profile of postorbital region of head uniformly plain but
gently bulged in cross-section, crests absent at postorbital region,
frontals developed and extended posteriorly; median dorsal-strut of
supraoccipital almost totally obliterated dorsally by the frontals, which
are in close proximity medially throughout their extent; a pair of slit-
like small fontanels present laterally to the supraoccipital median strut,
bordered anteriorly by the posterior margin of the frontals and poste-
riorly by the lateral walls of the supraoccipital; a median ridge on top
of head at orbital region separating the supraorbital canals laterally;
two bony bridges on each frontal between the median crest and the
lateral margin of the bone, shielding above the supraorbital canal; the
anterior bony bridge is located slightly anterior to the middle of the
orbit and is oriented at 90
in relation to the longitudinal axis of the
body; the posterior bony bridge is located slightly in advance of the
posterior margin of the orbit and is oriented at c. 45
in relation to the
longitudinal axis; anterior frontal fontanels absent, but bone at the
level of the anterior bridge somewhat soft and apparently able to sus-
tain some level of displacement; mesethmoid developed, compressed
laterally and projected anteriorly in relation to vomer, a small portion
10
FISH
of its posterodorsal region sandwiched between the anterior tips of
frontals at the top of the cranium; a pair of lateral processes on the
mesethmoid, the anterior one located at the anterior tip of the bone
and projecting laterally at approximately right angles to the longitudi-
nal axis of the body, the posterior one located in the posteroventral
region of the mesethmoid, close to the vomer, projected
posteroventrally and associated with the premaxilla; nasal and
antorbital extremely developed, forming an apparatus that supports
the greatly expanded nasal chamber.
Long and narrow gill rakers on first branchial arch, their oral sur-
faces densely covered with delicate teeth; total number of gill rakers
on first branchial arch 49(3), 50(7)*, 51(9), 52(13), 53(23), 54(22),
55(26), 56(16), 57(11), 58(12), 59(7), 60(1), 61(1); number of gill rakers
on lower portion of first branchial arch 28(2), 29(8), 30(33)*, 31(38),
32(30), 33(31), 34(7), 35(2); number of gill rakers on upper portion of
first branchial arch 19(1), 20(12)*, 21(21), 22(22), 23(30), 24(36),
25(19), 26(6), 27(2), 28(2); about six rakers also present at hind face of
the dorsal portion of third branchial arch; rakers absent from hind face
of the lower portion of third branchial arch and from both dorsal and
ventral portions of more anterior arches; basihyal cartilaginous,
reduced, but partially covered by a small tooth plate bearing a few
scattered delicate teeth (Figure 4a); an apparently single, narrow, con-
tinuous tooth plate with delicate teeth extending from the anterior
portion of basibranchial 1 to the middle ossified portion of
basibranchial 3; tooth plate over basibranchials not expanded laterally
over the pair of hypobranchial 2; posterior endochondral tip of
basibranchial 3 under anterior portion of the cartilaginous and rela-
tively long basibranchial 4; narrow delicate tooth plates densely
packed with minute teeth arranged in a series at the dorsal (oral) sur-
face of the lower portion of arches 1 and 2, including the respective
hypobranchials and ceratobranchials; a few scattered tooth plates
sometimes present on lower portion of third arch (i.e., ceratobranchial
3), not extensively covering the structure; tooth plates absent from
the lower portion of fourth arch (i.e., ceratobranchial 4); about 30 coni-
cal and developed teeth present on the posterior region of
ceratobranchial 5, fused with the underlying bone, teeth at the poste-
rior margin distinctly more developed than teeth at the anterior por-
tion of the bone; anterior elements of upper portion of branchial
arches strongly meeting in the midline (Figure 4c,d); gongyloid carti-
lage reduced, but clearly distinct (Figure 4d); small scattered tooth
plates present on oral (ventral) surface of upper gill arches, plates
associated with infrapharyngobranchials 2–3 fused with the underly-
ing bone; infrapharyngobranchial 3 cartilaginous, but two upper pha-
ryngeal tooth plates present and with well-developed dentition, the
more posterior in the shape of a narrow triangle, oriented post-
erolaterally; epibranchial 4 developed, but epibranchial organ appar-
ently absent or extremely reduced; branchiostegal rays 12 (5 C&S
specimens).
Dorsal-fin origin equidistant between anterior border of eye and
caudal-fin base; base of dorsal fin lined with small scales; dorsal-fin
rays ii(3) or iii(148)*, followed by 9(1), 10(3), 11(22), 12(79)*, 13(42) or
14(4) branched rays; longest dorsal-fin rays reaching to tip of last
branched dorsal-fin ray when fin depressed; pectoral-fin rays i(146)*,
AGOSTINHO ET AL.
followed by 10(1), 11(2), 12(23), 13(47), 14(69)* or 15(4); pectoral-fin
tip failing to reach pelvic-fin base by about the eye diameter, or
approximately 43%–68% (mean 50.8%) of pectoral-fin length in speci-
mens larger than 100 mm SL; axillary scale present, developed, almost
equal in length to pectoral fin; pelvic-fin origin slightly in advance of
middle distance between pectoral-fin origin and anal-fin origin; pelvic-
fin rays i(147)* or ii(1), followed by 5(3), 6(131)* or 7(14) branched
rays; last two pelvic-fin rays joined to each other and to the body by a
thin membrane for about half their length, a short triangular scale
between pelvic fins; axillary scale also present on pelvic fin, often lost
in specimens examined; pre-pelvic and post-pelvic scutes absent, a
single scute present anterior to pelvic-fin base (pelvic scute); median
keel of pelvic scute absent, ascending arms slightly inclined anteriorly,
their distal tips reaching about one-fourth of body height at that verti-
cal; anal-fin origin under or slightly behind vertical through the base of
last dorsal-fin ray, slightly nearer to the pelvic-fin origin than to
caudal-fin origin; base of caudal-fin lined with small scales; anal-fin
rays ii(2) or iii(146)*, followed by 11(11), 12(59), 13(71) or 14(7)*; cau-
dal fin markedly forked, upper and lower lobes about the same size; 1
(6) or 4(5) rudimentary fin rays (sensu Grande & Bemis, 1998) on upper
lobe of caudal fin, followed by iiii(150), 9(71), 10(63), 11(16) principal
(segmented) rays; 1(4) or 4(5) rudimentary fin rays on lower lobe,
followed by iiii(150), 9(65), 10(67), 11(18) principal (segmented) rays
(rudimentary caudal-fin rays counted only in C&S specimens); bases
of middle caudal-fin rays with a “two-peg” arrangement (sensu
Grande & Nelson, 1985), i.e., a single peg present on outer margin
of both rays, ventral peg absent in upper middle caudal-fin ray, dor-
sal peg absent in lower middle caudal-fin ray; a pair of developed
alar scales on each lobe of caudal fin, borders delicate and distinctly
emarginated in C&S specimens, condition not so evident in alcohol-
preserved specimens; outer alar scales more developed, the alar
scale associated with upper caudal-fin ray 5 (counting from the first,
middle, upper caudal-fin ray) gently curved downwards over the
scale associated with upper caudal-fin ray 4, the alar scale associ-
ated with lower caudal-fin ray 4 (counting from the first, middle,
ventral caudal-fin ray) gently curved upwards over the scale associ-
ated with lower caudal-fin ray 3; two epurals, two free uroneurals,
six independent hypurals; interlobar notch between hypurals 2 and
3 (Monod's hypural diastema) triangular in shape, somewhat
reduced (Figure 6); compound centrum at caudal skeleton formed
by the fusion of preural centrum 1, ural centrum 1, uroneural 1, and
hypural 2; interzygapophysal articulation developed, pre- and post-
zygapophyses particularly conspicuous between preural centrae 4–
17 (Figure 6).
Lateral line absent; scales cycloid, often lost; 41(8) or 42(12)
transverse series at the longitudinal line; anterior border of scales
with distinct “shoulders” and median prominence, unexposed portion
of scale with a single complete vertical striation, preceded in the
posterior scales by short radial striae; exposed portion of scale
with a vertical striation, often curved, the whole area reticulated
in posterior scales; scales along dorsal region with six to seven
longitudinal striae (most descriptive information of scales based on
Whitehead, 1973).
AGOSTINHO ET AL.
F I G U R E 6 Terminal portion of the vertebral column and caudal
skeleton of Anchoviella cayennensis (NPM 3265) in left lateral view.
Abbreviations: ep: epural; hyp1-6: hypurals 1–6; hs: haemal spine;
postz: neural and haemal postzygapophyses; prez: neural and haemal
prezygapophyses; pu1-4: preural centrum 1–4; u1-2: ural centrum 1–
2; un: uroneural. Scale bar = 2 mm
3.6 | Colouration in life
Most descriptive information is based on Puyo (1945, 1949). Body
light brown, small specimens probably translucent; opercular region
somewhat darker, with iridescent reflections, some areas with a
metallic blue hue; a well-marked silvery lateral stripe, slightly tapering
towards tail, a darker layer of melanophore stripe underneath the gua-
nine layer; posterior third of the body with diffuse transverse silvery
streaks projecting from the lateral stripe, directed towards the ventral
part of the caudal peduncle; irregularly shaped silvery spots also dis-
posed on the caudal-fin base; dorsal and anal fins hyaline to translu-
cent; pectoral and pelvic fins hyaline; caudal fin with a lighter
colouration compared with the body, upper and lower borders of the
caudal fin slightly tinged with brown.
3.7 | Colouration in alcohol
Body brown to yellowish; darker pigments more evident on specimens
collected in estuaries, oceanic specimens overall lighter in colouration;
silver to pale lateral stripe, often partially or totally absent in pre-
served specimens, when present extending from the posterior border
of the operculum to the caudal peduncle; darker line extending along
the dorsal margin of the silver stripe region present in most speci-
mens, even when silver stripe is absent due to guanine layer decay
after preservation; largest width of dorsal stripe 1–1.5 times orbit
diameter; black spots present along the dorsal profile, from the ante-
rior portion of head to the caudal fin, including dorsal-fin base; a dis-
tinct somewhat diamond-shaped blackish area present at nape region,
anterolaterally limited by the pair of posttemporal bones; top of head
in the region posterior to orbit also blackish, a median darker stripe
projecting anteriorly up to the tip of the nose; black spots also present
on anterior region of anal- and caudal-fin bases; iris and post-orbital,
FISH 11
gular and infraorbital regions with some guanine; fins hyaline but
some brownish to black dots on alar scales and along caudal-fin rays.
3.8 | Sexual dimorphism
Ripe females collected in estuaries and rivers seem to be overall larger
and more robust in terms of abdominal girth. However, those putative
differences were not recognized as valid according to statistical
analyses.
3.9 | Geographic distribution
Confirmed records of A. cayennensis in Museum Collections (Figure 7)
indicate the species is distributed in coastal and estuarine environ-
ments along the Atlantic coast of South America, from Suriname (Cop-
pename River near Hermina bank; North Atlantic) to the State of Rio

i; South Atlantic), southeastern Brazil.
de Janeiro (Itaipú beach, Nitero
More data are needed from northern Brazil, between the states of
Piauí and Pernambuco, where the species is also probably present.
Confirmed records of A. cayennensis and A. perfasciata in this study
suggest that the two species are not sympatric; however, additional
specimens are needed from northern South America.
3.10 | Ecological notes
Anchoviella cayennensis was described from two specimens collected
at the mouth of the Cayenne River, French Guiana. Puyo (1945)
reported additional specimens collected by fishermen in the Mahury
and Kourou Rivers, also coastal rivers in French Guyana. He appar-
ently regarded the species as not migratory, endemic to estuaries and
coastal rivers in the region [“on ne le pêche jamais en mer”; “Cette
espèce, à l'encontre des autres Stolephorus, doit être sédentaire”
(Puyo, 1945: 103)]. It is interesting that more than 70 years later
Barbosa et al. (2017) also considered A. sanfranciscana, now under the
synonymy of A. cayennensis, endemic to the estuary of the São Fran-
cisco River, northeastern Brazil. The fact that the type series of both
A. cayennensis and A. sanfranciscana was collected in estuaries might
not be just a coincidence; this relatively rare species of anchovy seems
to preferentially inhabit estuaries and lower portions of coastal rivers,
judging by the fact that most specimens examined in this study
(123 out of 151) were collected in those environments. However, the
species is at least occasionally collected in open marine coastal
waters. Valentim and Esteves (2018) reported that A. cayennensis
apparently migrate during spawning to the upper section of the lower
Paraíba do Sul River in southeastern Brazil. Still according to Valentim
and Esteves (2018), the reproductive peak of A. cayennensis in the
Paraíba do Sul River occurs during the dry season (March–October),
when artisanal catches increase significantly. Therefore, and contrary
to Puyo's (1945) initial assumption, it seems that A. cayennensis is a
migratory coastal species, perhaps with a preference for estuaries, but
12
FISH
more data on abundance, frequency and reproduction of the species
are needed, especially in large rivers such as the São Francisco, Ama-
zonas and Cayenne.
3.11 | Popular names
Cayenne anchovy (English), Manjuba, Manjubinha, Manjuba Perna-de-
Moça, Pilombeta, Pititinga (Brazilian Portuguese), Anchovieta de
Cayena (Spanish), Anchois de Cayenne, Jamais-gouté (French).
3.12 | Conservation status
Judging by the small size of lots examined (maximum of 23 specimens;
MZUSP 60368) and the relatively low total number of specimens
identified in ichthyological collections, A. cayennensis is apparently
naturally rare, especially when compared to more abundant sympatric
species of the genus, such as A. lepidentostole and A. guianensis. The
species is also now apparently rare in the estuary of the Cayenne
River (Tagliarolo, M. and Rousseau, Y. – “Institut Français de
Recherche pour l'Exploitation de la Mer – IFREMER Guiane,” pers.
comm., 2020). Lower portions of coastal rivers and estuaries are
F I G U R E 7 Localities of specimens of Anchoviella cayennensis
identified after the taxonomic revision (white circles: non-type
specimens; black circle: type locality), with additional locality
information from Whitehead (1973; red circles) and Hildebrand and
de Carvalho (1948; yellow circles)
AGOSTINHO ET AL.
highly impacted by several human activities, particularly in the São
Francisco River and in other rivers of the central coast of Brazil. The
species is now targeted along with A. lepidentostole in the Paraíba do
Sul River (Valentim & Esteves, 2018) and is also probably fished as by-
catch in artisanal fisheries throughout its distributional range. Despite
those possible impacts, the geographic distribution of A. cayennensis is
fairly broad, and at the moment there is no evidence indicating sub-
stantial declines in its population. Anchoviella cayennensis was recently
assessed as “least concern” by the IUCN at the global level
(Carpenter, 2015) and also in Brazil (ICMBio/MMA, 2018), which
includes most of its distribution, and we agree with those assessments
based on the new data uncovered here. Nonetheless, given both its
apparent rarity and likely dependence on estuaries and lower portions
of coastal rivers for reproduction, the species might be threatened
locally. Further studies are needed to properly identify and quantify
local and regional threats to the species.
F I G U R E 8 PCA of 22 morphometric characters with
191 specimens of Anchoviella examined in addition to the neotype of
Anchoviella cayennensis (MNHN-IC-2005-2320)
F I G U R E 9 Boxplot with the scores of each group in the first PCA
using the morphometric data presented in Table 1. (a, left) Anchoviella
cayennensis and the (a, right) type series of Anchoviella sanfranciscana
are not significantly distinct, but both groups are significantly distinct
from (b) Anchoviella perfasciata
AGOSTINHO ET AL.
FISH 13

3.13 | Morphological analyses
The PCA performed with all morphometric data (Figure 8) explains
46% of the total variation, with 33.1% explained by the first axis and
12.9% explained by the second axis. The first axis indicates the exis-
tence of two distinct groups, one formed by specimens identified as
A. cayennensis, including MNHN-IC 2005-2320 and the type series of
A. sanfranciscana, and the other formed by specimens identified as
A. perfasciata.
In the first PCA, 10 morphometric characters had a greater influ-
ence on the first axis in the separation of the two groups recovered:
HL, anal-fin base length, upper-jaw length, length of maxilla beyond
posterior margin of orbit, distance between the tip of pelvic-fin and
anal-fin origin, body height, interorbital distance, pre-anal length and
pelvic-fin length (Figure 8). Basically, the group formed by
A. cayennensis and the type series of A. sanfranciscana on the left of
PCA 1 is mostly characterized by specimens in which values of those
parameters are proportionately smaller in relation to SL and HL when
compared with specimens on the right, identified as A. perfasciata. A
boxplot with the scores of each of the groups (A. cayennensis,
A. perfasciata and the type series of A. sanfranciscana) in the first PCA,
using the morphometric data presented in Table 1, further supports
the conclusion that A. cayennensis and A. sanfranciscana are not
significantly different from each other but that both are distinct from
A. perfasciata (Figure 9).
As indicated in the Materials and Methods section, the second
PCA (Figure 10) was performed with the 10 measurements used by
Whitehead (1973) to redescribe A. cayennensis based on a single spec-
imen (RMNH 26294) collected at the mouth of the Coppename River
near Banco Hermina, Suriname. The second PCA explains 54.7% of
the total variation, with 37.9% on the first axis and 16.8% on the sec-
ond. The PCA again indicates the existence of two morphologically
distinct groups, one formed by specimens identified as A. cayennensis,
including RMNH 26294, in addition to the type series of
A. sanfranciscana, and the other formed by A. perfasciata.
In the second PCA, four morphometric characters were more rel-
evant to the ordination of specimens on the first axis: anal-fin base
length, upper-jaw length, HL and body height. In general terms, and
like the conclusions of PCA 1, those results indicate that the group
formed by A. cayennensis and the type series of A. sanfranciscana is
characterized by specimens in which values of those characters are
proportionately smaller in relation to SL and HL, when compared with
specimens of A. perfasciata.
The results of the analyses of morphometric data obtained in
both PCAs, therefore, confirm the existence of two morphologically
distinct groups among specimens examined, one of them including
A. cayennensis and A. sanfranciscana and the other including
A. perfasciata (Figures 8–10).
To test if the width of the silvery lateral stripe in relation to eye
diameter is a valid diagnostic feature of A. sanfranciscana, a linear

F I G U R E 1 0 PCA of 10 morphometric characters with F I G U R E 1 1 Linear regression of silvery lateral stripe width at
191 specimens of Anchoviella examined in this study and information dorsal-fin origin vs. eye diameter of the type series of Anchoviella
from Anchoviella cayennensis RMNH 26294 obtained from sanfranciscana (black circles) and 20 selected specimens of Anchoviella
Whitehead (1973) cayennensis (red circles)

T A B L E 3 Values (mm) of eye

A. cayennensis A. sanfranciscana
diameter and width of the silvery lateral
stripe at the dorsal-fin origin of 20 Selected 20 specimens Paratypes
selected specimens of Anchoviella
cayennensis and in the 20 specimens of Range Mean S.D. Holotype Range Mean S.D.

the type series of Anchoviella Eye diameter 7.5–9.2 8.3 0.7 7.5 7.2–8.4 7.8 0.4
sanfranciscana Width of silvery lateral stripe 7.2–12.2 10.1 1.1 8.6 8.5–10.4 9.1 0.6
14
FISH
T A B L E 4 Values (mm) of pectoral-fin length and distance between pectoral-fin tip and pelvic-fin base in 131 specimens of Anchoviella
cayennensis, including MNHN-IC 2005–2320, and in the type series of Anchoviella sanfranciscana
A. cayennensis
MNHN-IC 2005–2320
Pectoral-fin length 15.2
Distance between pectoral-fin tip and 4.7
pelvic-fin base
F I G U R E 1 2 Linear regression of pectoral-fin length vs. distance
between tip of pectoral-fin and base of pelvic fin of the type series of
Anchoviella sanfranciscana (black circles) and 131 specimens of
Anchoviella cayennensis (red circles)
regression was performed. In this analysis, values of the width of the
silvery stripe measured at the origin of the dorsal fin (where the
stripe is typically larger) and values of the eye diameter of the
20 specimens that compose the type series of A. sanfranciscana were
compared to the same features of 20 selected specimens of
A. cayennensis where the stripe was significantly intact after preser-
vation (Table 3). The linear regression based on values of eye diame-
ter vs. the width of the silvery lateral stripe again failed to
discriminate between the two sets of specimens; i.e., the width of
the silvery lateral stripe is not diagnostic of A. sanfranciscana
according to the results obtained (Figure 11).
Another diagnostic character of A. sanfranciscana proposed by
Barbosa et al. (2017) is the distance between the pectoral-fin tip and
the pelvic-fin origin similar to pectoral-fin length vs. the distance
between the pectoral-fin tip and the pelvic-fin origin less than half the
pectoral-fin length in A. cayennensis (Barbosa et al., 2017: 166). To
test the validity of this character as diagnostic between the two spe-
cies, another linear regression was performed with measurements of
those parameters in the type series of A. sanfranciscana and in
131 specimens of A. cayennensis (Table 4). The results of this linear
regression also failed to discriminate between the two sets of speci-
mens; that is, the distance from the pectoral-fin tip and pelvic-fin base
is not different in A. cayennensis compared with the type series of
A. sanfranciscana (Figure 12).
AGOSTINHO ET AL.
A. sanfranciscana
Other specimens Paratypes
Range Mean S.D. Holotype Range Mean S.D.
14.2–20.1 16.8 1.6 15.3 15–17 16 0.5
2.4–14 7.6 2.6 8.3 6.3–11.2 8.5 1.2
4 | DI SCU SSION
The results presented here clearly indicate that A. sanfranciscana is a
junior synonym of A. cayennensis. Anchoviella perfasciata, in turn, is a
valid distinct species. Anchoviella sanfranciscana was described based
on a small number of specimens (20) and with a limited number of
specimens of similar species in the comparative material (Barbosa
et al., 2017). The morphological similarity between A. sanfranciscana
and A. cayennensis was recognized by the authors during the descrip-
tion of the species. Basically, three characters were indicated as diag-
nostic of A. sanfranciscana by Barbosa et al. (2017): the presence of a
wide silvery lateral stripe, with a well-defined dark brown and yellow
lateral band above it, its total width 1.5–2 times wider than the ocular
(presumably eye) diameter; the distance from the tip of the pectoral fin
(apparently wrongly referred to as dorsal fin in the abstract and on page
164 of Barbosa et al., 2017) and the pelvic-fin base (apparently wrongly
referred to as the anal fin in the abstract and on page 164 of Barbosa
et al., 2017) similar in length to the pectoral-fin length; and the number
of gill rakers in the lower branch of the first branchial arch “unique,”
29–35.
Characters proposed by Barbosa et al. (2017) as diagnostic for
A. sanfranciscana are rather problematic or not clearly defined, per-
haps reflecting the absence of a larger number of comparative speci-
mens. The presence of a silvery lateral stripe, for instance, is shared
by several species of anchovies, but the stripe tends to reduce signifi-
cantly or totally disappear in most engraulids stored in scientific col-
lections. As noted earlier, the results of the current study indicate that
there is no significant difference between the width of the lateral
stripe when the type series of A. sanfranciscana is compared with
specimens of A. cayennensis. Also contrary to the observations of
Barbosa et al. (2017), the current analyses failed to indicate that the
distance between the tip of the pectoral-fin to the pelvic-fin origin is
different between those two sets of specimens.
Barbosa et al. (2017) further added that, despite all anatomical sim-
ilarities between A. sanfranciscana and A. cayennensis, A. sanfranciscana
has “a more robust body than A. cayennensis and is anatomically similar
to … A. lepidentostole” [misspelled as “lepisdostostole” in the original text
(Barbosa et al., 2017: 162)]. This supposedly more robust body shape
of A. sanfranciscana compared with that of A. cayennensis was also
suggested in X-ray pictures of specimens of the two species presented
by the authors (Barbosa et al., 2017: figure 10). Nonetheless, in addition
AGOSTINHO ET AL.
FISH 15

to this character not being clearly defined, morphometric data pres- misidentified as A. cayennensis in that region. In addition to the character-
ented here (Table 1) indicate that the overall body shape of the type istics indicated in the taxonomic key, A. cayennensis seems to attain a
series of A. sanfranciscana is not different from that of A. cayennensis larger SL when compared with A. guianensis (145.5 vs. c. 70 mm), a con-
and again likely results from the lack of more comparative material on dition that may help identify specimens.
the description of A. sanfranciscana.
Barbosa et al. (2017) also noted a similar value of gill rakers
1a. Anal-fin origin located at or posterior to the 2
between A. sanfranciscana and A. cayennensis among congeners, a sit- vertical through posterior margin of last dorsal-
uation that is indeed confirmed by the meristic data presented here fin ray
(Table 2). Other features of the branchial arch anatomy of one cleared 1b. Anal-fin origin anterior to the vertical through 5
and stained paratype of A. sanfranciscana examined (INPA 053280) posterior margin of last dorsal-fin ray
are indistinguishable from those of A. cayennensis and are in accor- 2a. Upper-jaw length beyond posterior margin of A. guianensis
dance with the description presented in the diagnosis provided earlier. orbit about up to 9% of HL
Branchial arch anatomy of A. perfasciata, in turn, is rather distinct from 2b. Upper-jaw length beyond posterior margin of 3
that of A. cayennensis, being characterized overall by a more devel- orbit more than 13% of HL

oped basibranchial dentition, with the tooth plate over basibranchials 3a. A total of 18–24 rakers on lower portion of first A. vaillanti
branchial arch; 17–22 branched anal-fin rays
1–3 wider and expanded posteriorly over hypobranchial 2 (Figure 4b).
In summary, all characters proposed as diagnostic for 3b. A total of 25–35 rakers on lower portion of 4
first branchial arch; 11–16 branched anal-fin
A. sanfranciscana are shared with A. cayennensis, and no further differ-
rays
ences between those species could be unravelled in this study. Ancho-
4a. A total of 28–35 rakers on lower portion of A. cayennensis
viella perfasciata, in turn, is clearly distinct in terms of both morphometric first branchial arch; 11–14 branched anal-fin
and meristic data, in addition to features of the branchial arch dentition. rays; upper jaw 56.1%–69.4% (mean 65.1%) of
HL; reduced basibranchial dentition, consisting
of a narrow median tooth plate over
basibranchials 1–3, not expanded posteriorly
5 | K E Y TO M A R I N E A N D E S T U A R I N E over hypobranchial 2
A T L A N T I C S P E C I E S OF A N C H O V I E L L A 4b. A total of 25–30 rakers on lower portion of A. perfasciata
first branchial arch; 14–16 branched anal-fin
The following identification key for eight coastal and estuarine species of rays; upper jaw 69.2%–77.3% (mean 74.1%) of
Anchoviella is proposed based on the examination of specimens by the HL; developed basibranchial dentition, tooth
plate over basibranchials 1–3 wide and
authors and information from the literature, including original descrip-
expanded posteriorly over hypobranchial 2
tions, taxonomic revisions and taxonomic guides, such as Hildebrand
5a. Upper jaw shorter than lower jaw; upper jaw Anchoviella
(1963), Whitehead (1973), Figueiredo and Menezes (1978), Whitehead 51%–62% of HL brevirostris
et al. (1988), Nizinski and Munroe (2003), Loeb (2012, 2013) and Loeb
5b. Upper jaw longer than lower jaw; upper jaw 6
and Figueiredo (2014). Anchoviella carrikeri, Anchoviella jamesi (Jordan & 63%–80% of HL
Seale, 1926), A. juruasanga and A. hernanni Loeb et al., 2018, are strictly 6a. Upper-jaw length beyond posterior margin of Anchoviella
freshwater species of the genus from the Amazon basin and also from orbit about up to 5.9% of SL manamensis
the Orinoco in the case of A. jamesi (Loeb, 2012, 2013; Loeb et al., 2018; 6b. Upper-jaw length beyond posterior margin of 7
Loeb & Figueiredo, 2014; Van Der Sleen & Bloom, 2018; Whitehead orbit more than 7.5% of SL
et al., 1988). Their occurrence in estuaries and coastal rivers of the Atlan- 7a. “Panamensis-type” sensory channels over Anchoviella
tic coast of South America is highly unlikely; therefore, those species are operculum (Whitehead et al., 1988: 323, figure elongata
a); 10–12 branched pectoral-fin rays
not included in the key presented. Anchoviella vaillanti is an apparently
7b. “Walkeri-type” sensory channels over A. lepidentostole
strictly freshwater species endemic to the São Francisco River, north-
operculum (Whitehead et al., 1988: 323, figure
eastern Brazil (Loeb & Figueiredo, 2014; Whitehead et al., 1988). b); 12–14 branched pectoral-fin rays
Records of the species in the estuary of that river require verification,
but the species is included in the following taxonomic key. Ongoing stud-

n, 1987) and A. blackburni
ies indicate that Anchoviella perezi (Cervigo
Hildebrand, 1943, which are sometimes recognized as valid (e.g.,
Kullander & Ferraris, 2003; Di Dario et al., 2017; Di Dario, 2018; Fricke 6 | C O M P A R A T I V E M A T E R I A L EX A M I N E D
et al., 2022), are actually junior synonyms of A. jamesi and
A. lepidentostole, respectively (Loeb et al., 2018). Therefore, those species 6.1 | A. perfasciata
are not included in the key. Judging by the number of incorrect identifi-
cations of lots examined in this study from collections made in estuaries Holotype: MCZ 17965 (79.4 mm SL), Cuba.
and coastal rivers of northern South America, it seems that specimens of Non-type specimens: Cuba: USNM 117685 (8, 69.4–77.0 mm SL),
the relatively more abundant A. guianensis (Eigenmann, 1912) are often Cuba, 12 February 1943; FMNH 37549 (4, 68.5–81.0 mm SL),
16
FISH
Atlantic Ocean, Havana, 23
120 2900 N 082
240 1000 W. USA, Florida:
UF 208245 (6, 42.8–64.0 mm SL), Virginia Key, Bear Cut, Marine Lab
beach, 25 October 1960; UF 220202 (20, 47.4–72.0 mm SL), Virginia
Key, Bear Cut, Marine Lab beach, 20 October 1954. Martinica: USNM
117685 (1, 48.6 mm SL), Tortugas, 20 June 1940.
AUTHOR CONTRIBUTIONS
F.D.D., L.S.A. and M.V.L. outlined the study; L.S.A. and M.V.L. gath-
ered morphometric and meristic information; L.S.A. performed statisti-
cal analyses; F.D.D. provided osteological descriptions; all authors
wrote and revised the manuscript.
ACKNOWLEDGEMEN TS
We thank Erica P. Caramaschi and Mateus O. de Paula (Department of
Ecology/UFRJ) for donating specimens of A. cayennensis from the
Paraíba do Sul River (RJ) to the last author in 2012, an act of scientific
generosity that eventually resulted in the development of this study.
We are especially grateful to Ana Bottallo (MZUSP) and Zouhaira Gabsi
(MNHN) for providing pictures of and crucial anatomical information on
the specimen of A. cayennensis deposited in the Muséum National
d'Histoire Naturelle, Paris, now elected as neotype of the species. Ana
C. Petry, Carlos C. Barboza (NUPEM/UFRJ) and Paulo C. Paiva
(Department of Zoology/UFRJ) helped in statistical analyses. Carlos
C. Barboza also provided insightful remarks on an earlier version of the
manuscript. Arthur B. Bauer (PPG-CiAC/UFRJ) helped in preparing the
maps. We also thank A. P. Pozzobon, M. M. Mincarone, P. A. Catelani
and G. Vinicius (NUPEM/UFRJ) for several contributions to the devel-
opment of this study. For the loan and donation of specimens and sup-
port to visits of one or more authors to their institutions, we thank
M. Sabaj (ANSP); M. Brito (MZFS); J. Joyeux (CIUFES); C. McMahan
(FMNH); L. R. Py-Daniel (INPA); Z. Gabsi (MNHN); C. R. Moreira, M. R.
Britto and D. Moraes (MNRJ); W. B. Wosiacki, I. Maschio, D. Coutinho
and A. Akama (MPEG); M. de Pinna, A. Datovo and O. Oyakawa
(MZUSP); R. Feeney (NHMLA); J. G. Mann (TUBRI); R. Robins (UF); M.
M. Rotundo (UNISANTA); and J. T. Willians, S. Jewett and R. Vari (in
memoriam) (USNM). Further information on the occurrence (or, in this
case, of the apparent absence of) A. cayennensis in the Cayenne River
was gently provided by M. Tagliarolo and Y. Rousseau (Ifremer). This
study was conducted as part of the MSc dissertation of the first author,

s-Graduação em Ciências
which was developed at the Programa de Po
Ambientais e Conservação (PPG-CiAC/UFRJ). As members of the MSc
Committee, V. Slobodian (UNB) and M. M. Mincarone (NUPEM/UFRJ)
provided several relevant comments and suggestions to the manuscript.
Financial support to the first author was provided through an MSc fel-
lowship of the Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES). Additional support was provided by the project

s-
FAPERJ no. 22/2016 – Apoio Emergencial ao Programa de Po
Graduação em Ciências Ambientais e Conservação (PPG-CiAC), um
PPG Emergente da UFRJ no Interior do Estado, funded by Fundação
Carlos Chagas Filho de Amparo à Pesquisa do Estado do Rio de Janeiro
– FAPERJ. Financial support was also provided by CNPq (Conselho

gico – PROTAX
Nacional de Desenvolvimento Científico e Tecnolo
443302/2020) to F.D.D.
AGOSTINHO ET AL.
FUNDING INF ORMATI ON
CAPES, FAPERJ and PROTAX/CNPq.
SIGNIF ICANCE STATEMENT
Anchoviella perfasciata, A. cayennensis and A. sanfranciscana are similar
Atlantic species of anchovies with a complex taxonomic history. Here,
the authors show that A. sanfranciscana is a junior synonym of
A. cayennensis and that A. perfasciata is a distinct species. Geographic
distributions of the species are more clearly defined. Anchovies are
relevant to fisheries and coastal marine environments. The results of
the study contribute to a better understanding of the diversity of the
group in the Atlantic.
OR CID
Fabio Di Dario https://orcid.org/0000-0001-9256-8485
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