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Kuayguara etymatee sp. nov., a New Genus
and Species of Artotrogidae (Copepoda:
Siphonostomatoida) with an Uncommonly
Atrophied Leg 1
Amilcar Farias1,* , Elizabeth G. Neves1, and Rodrigo Johnsson1
1Universidade Federal da Bahia, Instituto de Biologia, LABIMAR – Crustacea, Cnidaria e fauna associada. Avenida Milton Santos, s/n,
Campus Ondina, Bahia, Brazil. *Correspondence: E-mail: amilcaar@gmail.com (Farias)
E-mail: elizabethneves@gmail.com (Neves); rjohnsson@gmail.com (Johnsson)
urn:lsid:zoobank.org:pub:94E98B05-130F-4BF9-8BCB-903842E3280E
Received 22 September 2024 / Accepted 14 February 2025 / Published 13 May 2025
Communicated by Daniel Stec
Artotrogidae Brady, 1880 is a cosmopolitan family with 23 valid genera and 131 known species. However,
a considerable number of these species were subject of reexaminations and redescriptions recently.
With the crescent number of new species discovered, it is becoming possible to better understand their
boundaries. This study presents a new genus and species of Artotrogidae, recovered from unidentied
hosts in debris of benthonic samples from Todos-os-Santos Bay, northeastern coast of Brazil. Kuayguara
etymatee gen. et sp. nov. exhibits an underdeveloped rst leg, which possess an unsegmented protopod
and 1-segmented exopod, a unique set of morphological characteristics that dierentiates it from all other
genera of the family.
Key words: Invertebrate associate, Symbiotic copepods, Systematics, Uncommon leg, Taxonomy
Citation: Farias A, Neves EG, Johnsson R. 2025. Kuayguara etymatee sp. nov., a new genus and species of Artotrogidae (Copepoda:
Siphonostomatoida) with an uncommonly atrophied leg 1. Zool Stud 64:10. doi:10.6620/ZS.2025.64-10.
BACKGROUND
Artotrogidae Brady, 1880 is a cosmopolitan
family with species found associated with a wide
variety of invertebrate hosts, ranging from sponges to
urochordates. Even so, many species of the family were
recovered from mixed benthonic samples (Boxshall and
Halsey 2004; Ivanenko et al. 2018; Lee and Kim 2023).
Nowadays the family includes 23 valid genera and 131
known species (Walter and Boxshall 2025).
A significant part of the family species was
described between the second half of the nineteenth
and the early decades of the twentieth century (Boeck
1859; Thorell 1859; Brady and Roberston 1876; Brady
1880 1899 1910; Thomson 1883; Scott 1888 1898 1905
1912; Giesbrecht 1895 1899; Sars 1918; Hansen 1923;
Wilson 1923 1924), period known as the “golden age
of the copepodology” (Damkaer 2002). However, since
the 1960s, the genera and species described during this
period have been the target of extensive reexaminations
and redescriptions that, allied with new species recently
discovered, have expanded the understanding about
their boundaries (Eiselt 1961 1965; Stock 1965 1966;
McKinnon 1988; Kim 1996; Johnsson and Rocha
2002; Johnsson and Neves 2005; Lee and Kim 2023).
Nonetheless, a significant number of genera of the
family is remaining monospecic (Walter and Boxshall
2025).
Despite the great number of known species, only
five artotrogid species are recorded from the South
Atlantic Ocean: Bradypontius ancistronus Neves and
Johnsson, 2008; Cryptopontius aesthetascus Neves
and Johnsson, 2008; Cryptopontius expletus Neves and
Johnsson, 2008; Cryptopontius pentadikos Farias, Neves
Zoological Studies 64:10 (2025)
doi:10.6620/ZS.2025.64-10
1
© 2025 Academia Sinica, Taiwan
and Johnsson, 2020 e Cryptopontius phyllogorgius
Farias, Neves and Johnsson, 2020 (Neves and Johnsson
2008; Farias et al. 2020). Aiming to cover this
knowledge gap this study investigations bring to light
the description of a unique new artotrogid genus and
species with a unique underdeveloped leg 1.
MATERIALS AND METHODS
The copepods were found at seven different
locations of Todos-os-Santos Bay (Fig. 1). Samples
were collected on the submersed surface of pier decks
incrusted with a diverse benthonic community, such
as sponges, barnacles, mollusks, hydroids, bryozoans,
ascidians, and corals. The material was fixed in 10%
formalin for 48 hours and posteriorly preserved in 70%
ethanol.
The copepods were sorted out under a stereo
microscope Zeiss Stemi 508. Specimens were analyzed
immersed in glycerin and mounted on temporary slides
with adhesive plastic rings to avoid crushing (Kihara
and Rocha 2009). Examination, photos, measurements,
and drawings were made with the aid of a Zeiss Axio
Lab.A1 microscope equipped with a digital camera
AxioCam ERc 5S connected to an iPad (7th generation)
with Zeiss Labscope Software (version 4.0.2). The
holotype and allotype were dissected directly on a
Fig. 1. Collection sites: A) Bahia state highlighted on the South America continent; B) Todos-os-Santos Bay; C) Sample area of: Private pier on Bom
Jesus dos Passos Is. [4], Public pier on Bom Jesus dos Passos Is. [5], Private pier on Madre de Deus Is. [6], and, Private pier on Bimbarras Is. [7]; D)
Sample area of: Marina de Itaparica pier [1], Estação de Medidas Magnéticas de Itaparica pier [2], and, Public pier on Barra do Paraguaçu beach [3].
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permanent slide with CMC-9 mounting media for the
appendages analysis (Kihara and Rocha 2009). The
Illustrations were vectorized with CorelDRAW 2021
software (version 23.1.0.389).
For the leg armature formula, Roman numerals
represent spines and Arabic numerals represent setae
(Huys and Boxshall 1991). For antennule armature,
Roman numerals represent ancestral segments (Huys
and Boxshall 1991).
The type material analyzed was deposited in
the Crustacea Zoological Collection of the Museu de
História Natural da Bahia of the Universidade Federal
da Bahia, Brazil.
For Confocal Laser Scanning Microscopy, a female
paratype and a male paratype were stained with a 1:1
solution of Congo Red and Acid Fuchsin overnight,
following the procedure described by Corgosinho et al.
(2017). The specimens were examined using an Axio
Observer.Z1 equipped with LSM900; images were
obtained through an experiment of extended depth of
focus on ZEN 3.0 (blue edition) and the final images
were post-processed using the maximum projection
method, detailed acquisition information on table 1.
RESULTS
TAXONOMY
Order Siphonostomatoida Burmeister, 1835
Family Artotrogidae Brady, 1880
Kuayguara gen. nov.
urn:lsid:zoobank.org:act:1FC6D5B3-2D74-4273-B1D1-
6C065F51CC8F
Diagnosis: Artotrogidae. Body cyclopiform,
dorso-ventrally flattened. Cephalic shield ornamented
with scale-like structures showing integumental organs.
Prominent dorsal crest and rostrum. Radial bands on
the lateral margins of the cephalosome. Epimera of the
cephalosome, second, and third pedigerous somites
projected posteriorly. Medio-posterior margin of the
second and third pedigerous somites with posteriorly
elevated projections medially aligned. Urosome
5-segmented on female, genital double-somite with two
pairs of produced posteriorly projected epimera. Male
urosome 6-segmented, genital somite and first post-
genital somite with a pair of projected epimera produced
posteriorly. Both female and male with paired genital
apertures, equal sized and ventrally located. Female
antennule 8-segmented. Male antennule 9-segmented,
with additional aesthetascs, showing an incomplete
segmentation on the terminal segment; and signicantly
large spine on segment 7. Antennal exopod represented
by seta, and 2-segmented endopod. Maxillule bilobed.
Maxilliped 5-segmented. Leg 1 uncommonly reduced,
with single-segmented ramus. Leg 1, 3, and 4 with coxa
and basis fused. Leg 2 and 3 biramous and 3-segmented.
Leg 4 lacking endopod.
Type species: Kuayguara etymatee sp. nov. by
original designation.
Etymology: The genus name Kuayguara is a
junction of the words “Kûa” (= bay) and “-ygûara”
(= inhabitant), from old-Tupi language, referring to
the Tupinambá people, the ethnic group that inhabited
Todos-os-Santos Bay before the arrival of the
Portuguese colonizers.
Remarks: When redefining Artotrogidae, Eiselt
(1961) described, among other characteristics, the
legs 1 to 4 as being: “normal ausgebildet oder in
verschiedenem Ausbaße reduziert, P4 bis zu seinem
völligen Fehlen”, (normally developed or reduced
to various degrees, leg 4 to its complete absence). In
fact, the different levels of reduction patterns of leg
4 on Artotrogidae performed an important role in the
differentiation of the genera, splitting the family in
a few groups: (1) Leg 4 biramous with 3-segmented
exopod and endopod (Abyssopontius Stock, 1985;
Antarctopontius Eiselt, 1965; Artogordion Ivanenko,
Bandera and Conradi, 2018; Bradypontius Giesbrecht,
1895; Cribropontius Giesbrecht, 1899; Glannapontius
Table 1. Confocal laser scanning microscopy (CLSM) detailed acquisition settings
Figure Objective Pinhole Laser Wavelength Detection Wavelength Detector Gain
Fig. 5a N-Achroplan 5x/0.15 M27 0.96 AU / 35 µm 561 nm; 2.00 % 555–607 nm 840 V
0.84 AU / 35 µm 640 nm; 3.00 % 615–700 nm 877 V
Fig. 5b N-Achroplan 5x/0.15 M27 0.95 AU / 35 µm 561 nm; 1.00 % 568–621 nm 840 V
0.84 AU / 35 µm 640 nm; 5.00 % 594–700 nm 877 V
Fig. 6b Plan-Apochromat 10x/0.45 M27 1.46 AU / 35 µm 561 nm; 0.30 % 555–593 nm 840 V
1.26 AU / 35 µm 640 nm; 2.00% 605–700 nm 877 V
Fig. 6b Plan-Apochromat 10x/0.45 M27 1.46 AU / 35 µm 561 nm; 0.20 % 555–593 nm 840 V
1.26 AU / 35 µm 640 nm; 3.00% 605–700 nm 877 V
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Holmes, 1998; Myzopontius Giesbrecht, 1895;
Neobradypontius Eiselt, 1961, Neopontius Scott
T., 1898; and; Sestropontius Giesbrecht, 1899).
(2) Leg 4 biramous with 3-segmented exopod and
2-segmented endopod (Arctopontius Sars G.O., 1915
and Metapontius Hansen, 1923). (3) Leg 4 with
3-segmented exopod, without endopod (Ascidipontius
Kim IH, 1996; Chejupontius Lee J and Kim IH, 2023;
Cryptopontius Giesbrecht, 1899; Dyspontius Thorell,
1859; Pteropontius Giesbrecht, 1895; Pulicitrogus Kim
I.H., 1998; and; Sewellopontius Ummerkutty, 1966).
(4) Leg 4 reduced to a protopod (Pseudotrogus Eiselt,
1961). (5) Leg 4 absent (Glyptotrogus McKinnon, 1988;
Artotrogus Boeck, 1859; and; Tardotrogus Eiselt, 1961).
Kuayguara gen. nov. shares the uniramous
leg 4 with a 3-segmented exopod with the 7 genera
included in the third group, but these taxa can be
easily differentiated by the level of reduction patterns
found on their leg 1: Ascidipontius, Chejupontius,
Cryptopontius, Dyspontius, and Pulicitrogus exhibit a
biramous 3-segmented leg 1, with the exopod formula:
(I-0, 0-0; II, 3), (I-0; 0-1; II, I, 2 or II, I, 1), (I-1; I-1;
III, 4 or III, 5), (I-1; I-1 or 0-1; II, 5 or II,4), (I-0; 0-1;
II, 3), respectively, characterizing minor modications
in the pattern, showing only a few elemental reductions
(Thorell 1859; Giesbrecht 1899; Sars 1918; Kim 1996
1998 2016; Johnsson 2001; Farias et al. 2021; Lee
and Kim 2023). This is completely different from
Kuayguara gen. nov. which shows only one-segmented
ramus on leg 1.
Sewellopontius species can exhibit a moderate
reduction pattern, with leg 1 exopod 2- or 3-segmented,
with the formula (I-0 or 0-0; 0-1; II, 3) or (I-0; II, 3
or II, 4) (Ummerkutty 1966; Kim 1996; Lee and Kim
2023). Therefore, also diering from the one-segmented
ramus of leg 1 in the new genus.
The species included in the genus Pteropontius
exhibited the most reduced pattern known to the leg
1, with both exopod and endopod 2-segmented, with
the formula (0-1 or I-0; II, 3 or II, 4). This pattern has
been conrmed with the recent redescription of the type
species by Lee and Kim (2023) and differentiates it
from the new genus.
The species P. pediculus exhibits an exception
to this pattern, it was described based on a single
male specimen found in association with the coral
Echinopora lamellosa (Esper, 1791) on Mauritius Is.,
Stock (1966) also classied the specimen as “somewhat
aberrant”, alluding to the uncommon morphology of
its reduced rst leg, which exhibit a single segmented
ramus.
Considering the erection of the new genus, it
is possible to observe other differences between P.
pediculus from all other Pteropontius species, such
as the 2-segmented antennal endopod (instead of
1-segmented); leg 1 and leg 3 with coxa and basis fused
(instead of articulated). Thus, since these characteristics
previously mentioned, plus the leg morphology, are
diagnostic of the new genus, P. pediculus should be
moved to Kuayguara gen. nov., with a new combination
as Kuayguara pediculus (Stock, 1966) comb. nov.
Kuayguara etymatee gen. et sp. nov.
(Figs. 2–5)
urn:lsid:zoobank.org:act: 9015C9A2-67FE-4578-8B3B-
904D3023ABEA
Material examined: Holotype ♀ (UFBA4130),
dissected on permanent slide; allotype ♂ (UFBA4133),
dissected on permanent slide; from mixed benthonic
samples, collected at public pier on Barra do Paraguaçu
beach (Salinas da Margarida city – 12°50'25.0"S,
38°47'40.9"W). Paratypes: 1♂ (UFBA4131) and,
1♀ (UFBA4132) collected at public pier on Barra
do Paraguaçu beach (Salinas da Margarida city –
12°50'25.0"S, 38°47'40.9"W). 1♀ (UFBA4092) and,
1♀ (UFBA4093) collected at Marina de Itaparica
pier, Itaparica Is. (Itaparica city – 12°53'21.2"S,
38°41'04.2"W). 1♀ (UFBA4112) collected at a private
pier on Bom Jesus dos Passos Is. (Salvador city –
12°45'42.4"S, 38°38'09.5"W). 1♂ (UFBA4098), 1♀
(UFBA4099), 1♂ (UFBA4100), 1♀ (UFBA4101),
1♀ (UFBA4102), 1♀ (UFBA4103), and 2♀
(UFBA4134), collected at Estação de Medidas
Magnéticas de Itaparica pier, Itaparica Is. (Itaparica
city – 12°52'47.9"S, 38°41'10.1"W). 1♂ (UFBA4137)
and 1♂ (UFBA4138), collected at a private pier on
Bimbarras Is. (Madre de Deus city – 12°43'36.1"S,
38°38'02.7"W). 3♀ (UFBA4316), 1♂ (UFBA4317),
2♀ and 1♂ (UFBA4139), 2♀ (UFBA4187), 1♀
(UFBA4353), 1♀ (UFBA4827), 4♀ (UFBA4826),
and 1♂ (UFBA4827) collected at a private pier on
Madre de Deus Is. (Madre de Deus city – 12°43'58.8"S,
38°37'22.8"W). 1♀(UFBA4777), 1♀ (4778), and a
juvenile (UFBA4779) collected at a public pier on Bom
Jesus dos Passos Is. (Salvador city – 12°45'20.7"S,
38°38'21.5"W). All paratypes from mixed benthonic
samples, preserved in ethanol.
Description of the female holotype: Body (Fig.
5a) cyclopiform, dorso-ventrally flattened, 888 µm
long. Cephalic shield adorned with scale-like structures
showing sensilla on each integumental organs (Fig. 5a,
b), and radial bands along all margins (Fig. 2b); body
length: width ratio 1.6:1. Cephalosome slightly wider
than long (539 × 553 µm); with robust dorsal crest,
ranging from posterior medial margin to prominent
rostrum; also bearing radial bands along margins
and ending in projected epimera. Second and third
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Fig. 2. Kuayguara etymatee sp. nov., Holotype female (UFBA4130): a, Body, dorsal view; b, radial bands and detailed ornamentation of the
carapace; c, urosome; d, antennule; e, antenna; f, oral cone and stylet tip; g, maxillule. Scale bars: a–b = 100 µm; c–e = 50 µm; f = 20 µm; g = 50 µm.
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pedigerous somites with serrulated margins, and
showing medial posterior margins with dorsal elevated
projections aligned with dorsal crest. Second pedigerous
somite significantly wider than long (88 × 327 µm),
with produced projected epimera. Third pedigerous
somite wider than long (100 × 287 µm), with elevated
structure on posterior medial margin projecting over
fth pedigerous somite. Fourth pedigerous somite twice
as wide as long (31 × 62 µm), and strongly reduced.
Prosome longer than wide (703 × 553 µm), prosome:
urosome length ratio 1.2:1.
Urosome (Fig. 2c) 5-segmented, longer than wide
(245 × 212 µm). Fifth pedigerous somite wider than
fourth one (30 × 96 µm). Genital double-somite nearly
twice as wide as long (110 × 214 µm) exhibiting two
pairs of projected epimera. Paired genital apertures on
the proximal region of the somite, equal sized, well-
spaced and ventrally located. Egg sac not observed. First
and second post-genital somites wider than long (28 ×
78 µm, and 19 × 56 µm). First somite partially covered
by genital one. Robust anal somite, wider than long (42
× 77 µm), showing row of five parallel perforations.
Caudal rami longer than wide (49 × 28 µm), armed with
two smooth and small dorsal setae, and distally armed
with 4 setae. Caudal rami length: width ratio 1.7:1.
Antennule (Fig. 2d) 8-segmented, total length
measuring 249 µm long. Length measurements made
along medial margin = 51, 63, 25, 18, 20, 12, 19, and,
41 µm long, respectively. Segmental homologies and
armature as follow: I – 1; II-VIII – 6; IX-XII – 4; XIII-
XIV – 1+s; XV-XVI – 1; XVII-XVIII – 1; XIX-XX – 1;
XXI-XXVIII – 10+ae. All setae naked, two distal long
setae almost as long as aesthetasc, which is 93 µm long.
Antenna (Fig. 2e) 4-segmented, total length
measuring 100 µm long (without distal setae), Coxa
and basis unarmed, 10 and 35 µm long, respectively.
Exopod reduced to seta, 22 µm long, almost as long as
first endopodal segment. Endopod 2-segmented, first
segment 25 µm long and unarmed, second segment
30 µm long, 1.2 times longer than first one, armed
medially with plumose seta, and distally with 3 setae:
a small slender seta, 2 stout ones, unequally sized, and
elongated plumose seta.
Oral cone (Fig. 2a) 260 µm long, reaching basis
of maxilliped. Mandible with long stylet inserted in oral
cone, showing 4 teeth-like projections at the tip, palp
absent (Fig. 2f).
Maxillule (Fig. 2g) bilobed, inner lobe longer
than outer one, 109 µm long (broken, as long as the
oral cone on paratypes), exhibiting a tapering shape
and naked margins, with long plumose seta; outer lobe
55 µm long, with row of setules on outer margin, and
two strong distal setae; longest seta has plumose inner
margin and spinules along outer margin, shortest one
shows spinules on inner margin and naked outer margin.
Maxilla (Fig. 3a) stout, syncoxa naked, measuring
202 µm long, with prominent bump on outer margin;
(detached) strong basis (claw), 230 µm long, remarkably
recurved, and armed with small stout seta on medio-
distal margin. Maxilliped (Fig. 3b) 5-segmented, total
length 291 µm long (not including distal claw); coxa 36
µm long, armed with single seta on inner margin, and
row of setules on outer margin; basis unarmed, 136 µm
long. Endopod 3-segmented, rst segment unarmed, 28
µm long; second segment 38 µm long, armed with distal
inner seta; third segment 53 µm long, armed with robust
seta and terminal claw. Terminal curved claw measuring
82 µm long.
Leg 1 (Fig. 3c) showing no segmentation between
coxa and basis, forming protopod, and exhibiting long
seta on distal outer margin. Leg 1 (Fig. 3c) has a single-
segmented ramus, armed with 3 distal unequal-sized
setae.
Leg 1 to 4 armature formula as follows:
Coxa Basis Exopod Endopod
Leg 1 - - 3 -
Leg 2 0–1 1–0 I-1, I-1, III, I, 5 0–1, 0–2, 1,2,3
Leg 3 0–0 1–0 I-1, I-1, III, I, 5 0–1, 0–2, 1, 1+I, 3
Leg 4 0–0 1–0 I-1, I-1, III, I, 4 absent
Outer margins of exopodal segments of legs 2 to
4 (Fig. 3d–f) armed with spinules. Endopodal segments
of leg 2 (Fig. 3d) showing setules along outer margin.
Third endopodal segment of leg 3 (Fig. 3e) exhibiting
strong distal spine. Distal spine of the third exopodal
segment of legs 3 and 4 (Fig. 3e–f) slightly bending on
the proximal region.
Both protopod and exopod of leg 5 (Fig. 3g)
extremely reduced, represented by 1 and 2 setae,
respectively.
Description of the male allotype: Body (Fig.
4a) cyclopiform, longer than wide, smaller but
comparatively slightly slender than female (530 ×
292 µm), body length: width ratio 1.8:1. Body adorned
as seen on female (Fig. 6a, b). Cephalosome longer
than wide (319 × 292 µm), cephalosome length:
width ratio: 1.1:1. Prosome longer than wide (402
× 292 µm), prosome: urosome length ratio 2.5:1.
Urosome 6-segmented, longer than wide (158 ×
110 µm). Genital somite (Fig. 4b) wider than long (63
× 148 µm), with only a pair of epimera posteriorly
projected. Paired genital apertures, well-developed,
equal-sized, and ventrally located. First post-genital
somite almost as wide as genital somite (35 × 123 µm),
also showing epimera posteriorly projected. Second
post-genital somite almost entirely covered by rst one,
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Fig. 3. Kuayguara etymatee sp. nov., Holotype female (UFBA4130): a, maxilla; b, maxilliped; c, leg 1; d, leg 2; e, leg 3; f, leg 4; g, leg 5. Scale bars: a–
f = 50 µm; g = 20 µm.
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Zoological Studies 64:10 (2025)
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Fig. 4. Kuayguara etymatee sp. nov., Allotype male (UFBA4133): a, body, dorsal view; b, urosome; c, antennule; d, maxilla; e, leg 1; f, third
exopodal segment of leg 3. Scale bars: a = 100 µm; b–e = 50 µm.
page 8 of 14Zoological Studies 64:10 (2025)
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and followed by third somite, both wider than long (17
× 63 µm and 13 × 48 µm), respectively. Anal somite
robust, almost twice wider than long (36 × 63 µm).
Caudal rami longer than wide (31 × 22 µm), bearing
two dorsal small naked setae and four terminal setae,
caudal rami length: width ratio 1.4:1.
Antennule (Fig. 4c) 9-segmented, total length
along medial margin measuring 312 µm long. Segments
measurements: 31, 60, 12, 15, 35, 23, 49, 27, and,
60 µm long, respectively. Segmental homologies as
follows: I – 0; II-VI – 3+5ae; VII – 1+ae; VIII – 0; IX-
XII – 2+s+3ae; XIII-XIV – 1+ae; XV-XVIII – 3+s+ae;
XIX-XX – 1+ae; XXI-XXVIII – 9+ae. All setae naked.
Large spine on segment XV-XVIII measuring 46 µm
long, almost reaching proximal margin of the last
segment. Terminal aesthetasc measuring 147 µm long.
Maxilla (Fig. 4d) with unarmed syncoxa, 179 µm
long, narrower than in female; and strong basis (grasping
claw) 208 µm long, armed with small stout seta on
medio-distal margin.
First leg (Fig. 4e) sharing female uncommon
reduced morphology. Protopodal segment bulkier
(65 µm long × 49 µm wide), exhibiting small seta on
distal inner margin in addition to long seta on distal
outer margin. And finally, third leg (Fig. 4f) showing
small distal seta on third exopodal segment.
All other features as seen on female.
Etymology: The species name etymatee is a
junction on the words “etymã” (= leg) plus “teé” (= very
dierent), both from old-Tupi language, alluding to the
uncommon morphology of leg 1 of the new species.
Remarks: The only congener of the new species,
Kuayguara pediculus (Stock, 1966) comb. nov., was
described based on a male, and some sexual dimorphic
features can only be compared with the male of the new
species (Table 2). The male body from both species
is similar, Kuayguara etymatee gen. et sp. nov. diers
from K. pediculus comb. nov. based on features such as
the circular-shaped radial bands on the lateral margins
of the cephalosome (instead of elongated); the medio-
posterior projection of the third pedigerous somite not
only covering the fifth but also covering the proximal
part of the genital somite (instead of not reaching the
genital somite); and finally, the second post-genital
somite not exhibiting posteriorly projected epimera
(instead of the projected epimera observed on K.
pediculus comb. nov. (Stock 1966).
The antennule of the new species exhibits no
aesthetasc on segment 4, three on segment 5, none on
segment 7; and the terminal aesthetasc is only twice
thicker than the others. While the male antennule of K.
pediculus comb. nov. shows an aesthetasc on segment 4,
two on segment 5, two on segment 7; and the terminal
aesthetasc is 4 times thicker than the others (Stock
1966).
The antenna Kuayguara etymatee gen. et sp.
nov. exhibits a naked exopodal seta, and unadorned
endopodal margins. While the male antenna of K.
pediculus comb. nov. shows features not seen in the
new species, such as a plumose exopodal seta, a row of
setules on the rst endopodal segment, and the crest-like
structures on the inner margin of the first and second
endopodal segments (Stock 1966).
The maxillule of K. pediculus comb. nov. shows
a row of setules on the proximal inner margin, and a
minute lateral seta on the distal margin of the inner lobe
(Stock 1966); both features not observed in the new
species.
The maxilla basis (grasping claw) of Kuayguara
etymatee gen. et sp. nov. exhibits only one stout seta;
while K. pediculus comb. nov. shows 3 spine-like
projections and a row of denticles (Stock 1966).
The maxilliped of the new species shows a naked
seta on the first segment, a smooth second segment,
Table 2. Comparison between Kuayguara gen. nov. species
Kuayguara etymatee gen. et sp. nov. Kuayguara pediculus (Stock, 1966) comb. nov.
Radial bands shape circular-shaped elongated
Dorsal medio-posterior projection of the 3rd ped.
somite
covers the proximal part of the genital somite did not reach the genital somite
Antennule terminal aesthetask 2x thicker than others 4x thicker than others
Antenna exopod seta naked plumose
Antenna endopod margins unadorned adorned with setules and crest-like structures
Maxillule inner lobe inner margin unadorned adorned with setules and minute seta
Maxilla basis stout seta 3 spine-like projections and denticles
Maxilliped armature formula (0, 1, 1) (2, 1, 1)
Leg 1 exopod smooth margins adorned with setules
Leg 3 third exopodal segment armature formula (1, 1+I, 3) (1, 2, 3)
Leg 5 seta 2 setae 3 setae
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© 2025 Academia Sinica, Taiwan
and endopodal armature formula (0, 1, 1); while K.
pediculus comb. nov. shows a plumose basal inner
seta on the rst segment; two rows of long setules, and
a medial serrated spine on the second segment; and
endopodal armature formula (2, 1, 1) (Stock 1966).
Despite sharing the uncommon morphology of the
legs, such as the reduction of the first, and the fusion
of the coxa and basis on the third and fourth legs;
Kuayguara etymatee gen. et sp. nov. exhibits a few
more dierences to point out from K. pediculus comb.
nov.: on the rst leg, the single-segmented ramus shows
smooth margins (instead of row of setules on outer
margins); the armature formula of the third endopodal
segment of leg 3 is (1, 1+I, 3) (instead of (1, 2, 3)); and
the fifth leg exopod is represented by 2 (instead of 3)
terminal setae (Stock 1966).
Key to Artotrogidae genera (adapted from
Boxshall and Halsey 2004)
1. L4 biramous ................................................................................ 2
- L4 uniramous, represented by protopod, or absent .................. 13
2. Epimeral plates of third pedigerous somite not produced beyond
anterior margin of genital double-somite ................................... 3
- Epimeral plates of third pedigerous somite produced with
posterolateral lobes extending posteriorly to anterior margin of
double-somite ............................................................................. 9
3. Female antennule 14-segmented; leg 4 endopod 2-segmented .....
............................................................ Metapontius Hansen, 1923
- Female antennule 9 to 14-segmented; leg 4 endopod 3-segmented
..................................................................................................... 4
4. Leg 5 reduced to single seta; antennary exopod absent, unarmed
............................................................ Abyssopontius Stock, 1985
- Leg 5 with at least 1 free segment (exopod) bearing 3 to 5 setae;
antennary exopod with free segment bearing 2 setae or without
free segment and represented by 2 setae .................................... 5
5. Antennary exopod 1-segmented ................................................. 6
- Antennary exopodal segment not expressed, represented by 2
setae on basis ................................. Glannapontius Holmes, 1998
6. Antennary exopod 1-segmented, armed with 3 setae; leg 5 armed
with 2 tooth-like processes ............................................................
........................ Artogordion Ivanenko, Bandera & Conradi, 2018
- Antennary exopod 1-segmented, armed with 2 setae; leg 5
without tooth-like processes ....................................................... 7
7. Leg 5 with free segment bearing 5 setae; antennary exopod with
2 setae ............................................. Antarctopontius Eiselt, 1965
- Leg 5 with free segment bearing 3 setae; antennary exopod with
2 setae ......................................................................................... 8
8. Oral cone with long distal siphon; maxillule outer lobe longer
than inner lobe ............................. Myzopontius Giesbrecht, 1895
Fig. 5. Confocal Laser Microscopy. Kuayguara etymatee sp. nov., Paratype female (UFBA4353) a, body, dorsal view; b, body, ventral view. Scale
bars: a–b = 200 µm.
page 10 of 14Zoological Studies 64:10 (2025)
© 2025 Academia Sinica, Taiwan
- Oral cone short, pear-shaped, without siphon; maxillule outer
lobe shorter than inner lobe .................. Neopontius Scott T, 1898
9. Epimeral plates of third pedigerous somite reaching mid-region
of genital double-somite ........................................................... 10
- Epimeral plates of third pedigerous somite extending beyond
genital double-somite and reaching second free abdominal somite
........................................................ Neobradypontius Eiselt, 1961
10. Endopod and exopod of leg 4 with similar length ........................
..................................................... Sestropontius Giesbrecht, 1899
- Endopod of leg 4 shorter than exopod ..................................... 11
11. Posterolateral angle of dorsal cephalic shield bid .......................
.................................................... Cribropontius Giesbrecht, 1899
- Posterolateral angle of dorsal cephalic shield simple ............... 12
12. Leg 1 with 3 outer spines on exopodal segment 3; leg 4 with
3-segmented endopod ................. Bradypontius Giesbrecht, 1895
- Leg 1 with 2 outer spines on exopodal segment 3; leg 4 with
2-segmented endopod ...................... Arctopontius Sars GO, 1915
13. Leg 4 absent ............................................................................. 21
- Leg 4 present ............................................................................ 14
14. Leg 4 reduced to protopod bearing 1 or 2 setae on process at
outer distal angle ................................ Pseudotrogus Eiselt, 1961
- Leg 4 uniramous, lacking endopod (or rarely biramous with
endopod represented by a single seta on minute segment as seen
in Cryptopontius digitatus Kim IH, 1996) ............................... 15
15. Leg 1 showing some level of segmentation reduction ............. 16
- Both rami of leg 1 3-segmented ............................................... 18
16. Leg 1 biramous .......................................................................... 17
- Leg 1 uniramous, exopod 1-segmented ....... Kuayguara gen. nov.
17. Both rami of leg 1 2-segmented ....................................................
...................................................... Pteropontius Giesbrecht, 1895
- Exopod of leg 1 2- or 3-segmented, endopod 3-segmented ..........
................................................ Sewellopontius Ummerkutty, 1966
18. Leg 1 with 3 outer spines and 4 or 5 setae (III,4/5) on third
exopodal segment ...................... Cryptopontius Giesbrecht, 1899
- Leg 1 with 2 outer spines on third exopodal segment .............. 19
19. Leg 1 with 2 inner setae on second endopodal segment .......... 20
- Leg 1 with 1 inner seta on second endopodal segment ............ 21
20. Leg 1 with 2 spines and 4 setae (II,4) on third exopodal segment
............................................................... Dyspontius Thorell, 1859
- Leg 1 with 2 spines and 3 setae (II, 3) on third exopodal segment
............................................................ Pulicitrogus Kim IH, 1998
21. Second exopodal segment of leg 1 unarmed .................................
.......................................................... Ascidipontius Kim IH, 1996
- Second exopodal segment of leg 1 armed with inner seta ............
............................................. Chejupontius Lee J & Kim IH, 2023
22. Exopod of antenna 1-segmented, bearing 3 setae .........................
...................................................... Glyptotrogus McKinnon, 1988
- Exopod of antenna represented by single seta on small papilla ....
................................................................................................... 23
23. Leg 1 with 2 inner setae on second endopodal segment and
typically with 3 outer spines on third exopodal segment ..............
................................................................. Artotrogus Boeck, 1860
- Leg 1 with 1 inner seta on second endopodal segment and
typically with 2 outer spines on third exopodal segment ..............
................................................................ Tardotrogus Eiselt, 1961
Fig. 6. Confocal Laser Microscopy. Kuayguara etymatee sp. nov., Paratype male (UFBA4827) a, body, dorsal view; b, body, ventral view. Scale
bars: a–b = 100 µm.
page 11 of 14Zoological Studies 64:10 (2025)
© 2025 Academia Sinica, Taiwan
DISCUSSION
The discovery of new taxa has allowed the
unraveling of known and until then unnoticed
taxonomical problems, as an example, the description
of the Neobradypontius Eiselt, 1961 and Metapontius
Hansen, 1923 that allowed Eiselt to propose that
the Artotrogidae family should be merged with
Dyspontiidae and Myzopontiidae (Eiselt, 1961). Most
recently, in a study with artotrogid species from Korean
waters, Lee and Kim (2023) described a new genus
and 7 new species for the family; among them, two
new species to the genus Sewellopontius, until then
monospecic (Lee and Kim 2023). With the knowledge
about the newly discovered Sewellopontius species
it was possible to observe that Pteropontius trimerus
Kim IH, 1996 exhibited features more related to the
latter, which resulted in the combination novae for the
species, as Sewellopontius trimerus (Kim IH, 1996)
(Lee and Kim 2023). The same process occurred with
the combination novae proposed in this study, with the
erection of the new genus it was possible to observe
that K. pediculus comb. nov. exhibits many dierences
from other Pteropontius species, and shares diagnostic
features with the new taxon.
The variation in leg armature formula is a crucial
systematic tool to understand the diversity of the
associated copepods, from species to ordinal level.
Moreover, fusions or losses of ramal segments can be
observed in most copepod orders, being one, both rami,
or the entire leg absent (Huys and Boxshall 1991). The
uncommon morphology of the first leg exhibited by
Kuayguara gen. nov. diverged from the most recent
artotrogid diagnosis, which states that “legs 1 to 4
biramous, typically with 3-segmented rami except
endopod of leg 4 reduced or absent; leg 4 entirely
absent in some genera. Leg 1 sometimes with reduced
segmentation in one or both rami, as in Pteropontius”
(Boxshall and Halsey 2004, p. 710). Consequently, the
diagnosis of the family should be extended to include
the possibility of a single-segmented ramus. The first
leg shall be diagnosed as “typically biramous and
3-segmented, sometimes with reduced segmentation in
one or both rami in some genera, exhibiting 2-segmented
rami or a single-segmented ramus”.
Contrarily to the observed on the extended
diagnosis of Entomolepididae in Farias et al. (2021),
in which the inclusion of a newly discovered biramous
and 3-segmented leg 4 possibly approaches the
family with Asterocheridae, the changes reported on
the Artotrogidae diagnosis brought new evidences
to distance it from its sister family. The uncommon
morphology of leg 1 on the new Artotrogidae genus
indicates a transformation towards reduction of the
limb, characterizing an apomorphy.
The study of the development of the legs of
Dioithona oculate (Farran, 1913) made by Ferrari
(1998) may help understand the odd morphology of
leg 1 exhibited by Kuayguara gen. nov. species. On
this investigation, five developmental stages were
described, corresponding to copepodid I to V. In the
first stage, the leg exhibited a bilobed bud with 3 and
2 setae, presumably indicating the initial formation of
the exopod and endopod, respectively. In the second
stage, the segment bearing the setae is transformed
on a biramous limb. And posteriorly, during the next
stages, new segments are added proximally on the
distal segment forming the 3-segmented biramous leg
(Ferrari 1998). The morphology of leg 1 of Kuayguara
gen. nov. suggests that the development of the limb
was suppressed in the first stage, not undergoing the
transformation expected to appear on copepodid II.
Despite the comprehension of the development
process involved on the leg 1 formation of the
new genus, its functional implications may also
be considered. On a study about copepods from
hydrothermal ecosystems, Heptner and Ivanenko (2002)
made a functional analysis on the morphology of
locomotory limbs; characterizing swimming, crawling,
and swimming-crawling types of copepods. Features
used on the characterization of crawling copepods are
the fused coxa and basis [1] (legs 1, 3, and 4) forming a
wide protopod [2] (legs 3 and 4), the way leg branches
are positioned folded up [3] to the flattened body [4].
These characteristics are observed on Kuayguara gen.
nov. While the presence of intercoxal plates [1] (legs
1, 2, and 3), the articulated and similar-sized coxa and
basis (leg 2) [2], and the swimming seta on all legs
[3] are features described as found on the swimming
type copepods. By exhibiting traits of both types,
Kuayguara gen. nov. features suggest the intermediate
characteristics of the swimming-crawling type of
copepods.
Ultimately, among the siphonostomatoid species,
there are some highly modied families in which one or
many legs are absent; but only Nanaspididae Humes and
Cressey, 1959 (holothurians parasites); Pseudocycnidae
Wilson, 1922 (fish parasites); and Stellicomitidae
Humes and Cressey, 1958 (starsh parasites) species are
recorded to have a reduced biramous and 1-segmented
leg 1 (Boxshall and Halsey 2004). Thus, leg 1 with a
single-segmented ramus of Kuayguara etymatee gen. et
sp. nov. is so far a unique feature on Siphonostomatoida.
page 12 of 14Zoological Studies 64:10 (2025)
© 2025 Academia Sinica, Taiwan
CONCLUSIONS
A new artotrogid genus and species, Kuayguara
etymatee sp. nov., is described from Todos-os-Santos
Bay, Northeastern Brazil, from mixed benthonic
samples. The new genus is the first artotrogid genus
to be described for Southwestern Atlantic waters,
and add up the number of Artotrogidae species
registered on the Brazilian coast to 6. In this study,
Confocal Laser Microscopy is used for the first time
in the family, showing details of some morphological
traits, as the first leg exhibiting a single-segmented
ramus, an unprecedented leg morphology on the order
Siphonostomatoida.
Acknowledgments: AF thanks the LABIMAR
members for the logistical support, and for the
efforts during sampling and sorting. The authors are
grateful to the Chico Mendes Institute for Biodiversity
Conservation (ICMbio) for collecting permission
(Sisbio No 15161-1). This study is part of the project
‘Assessment and research of sun coral in Todos-os-
Santos Bay’, a cooperation agreement between UFBA
and CENPES/PETROBRAS. This work was supported
by the CENPES/PETROBRAS [under grant number
5850.0107361.18.9].
Authors’ contributions: AF: collects, investigation,
morphological description, illustrations, confocal
analysis, and writing. EN: funding acquisition, review.
RJ: funding acquisition, review, supervision. All authors
read and approved the nal manuscript.
Competing interests: Authors declare no competing
interests.
Availability of data and materials: All data are
available in the paper. Type specimens were deposited
in the Crustacea zoological collection of the Museu de
História Natural da Bahia of the Universidade Federal
da Bahia, Brazil.
Consent for publication: Not applicable.
Ethics approval consent to participate: Not
applicable.
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