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A New Species of Snapper Perciformes Lutjanidae FR PDF
A New Species of Snapper Perciformes Lutjanidae FR PDF
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Abstract
Snappers of the family Lutjanidae contain several of the most important reef-fishery species in the tropical western
Atlantic. Despite their importance, substantial gaps exist for both systematic and ecological information, especially for
the southwestern Atlantic. Recent collecting efforts along the coast of Brazil have resulted in the discovery of many new
reef-fish species, including commercially important parrotfishes (Scaridae) and grunts (Haemulidae). Based on field col-
lecting, museum specimens, and literature records, we describe a new species of snapper, Lutjanus alexandrei, which is
apparently endemic to the Brazilian coast. The newly settled and early juvenile life stages are also described. This spe-
cies is common in many Brazilian reef and coastal estuarine systems where it has been often misidentified as the gray
snapper, Lutjanus griseus, or the schoolmaster, L. apodus. Identification of the new species cast doubt on prior distribu-
tional assumptions about the southern ranges of L. griseus and L. apodus, and subsequent field and museum work con-
firmed that those species are not reliably recorded in Brazil. The taxonomic status of two Brazilian species previously
referred to Lutjanus, Bodianus aya and Genyoroge canina, is reviewed to determine the number of valid Lutjanus species
occurring in Brazil. Twelve species of Lutjanus are now recognized in the western Atlantic, eight of which occur in Bra-
zil (one endemic). A key for the identification of all western Atlantic Lutjanus species and their known distributional
ranges is also provided.
Introduction
The fish fauna of the tropical western Atlantic is relatively homogeneous, but recent evidence demonstrates
that the southwestern Atlantic holds a considerable number of endemic species (e.g., Menezes et al. 1997,
2003). As a result of major collection-building efforts in the last decade, several groups of demersal fishes
have been investigated in the southwestern Atlantic, resulting in the identification of new species of myxinids
(Mincarone 2000, 2001), elasmobranchs (Gomes et al. 2000, Gomes & Paragó 2001), batrachoidids (Menezes
& Figueiredo 1998), atherinids (Bemvenuti 1995), grammids (Sazima et al. 1998), haemulids (Rocha & Rosa
1999), chaetodontids (Burgess 2001), pomacentrids (Moura 1995, Gasparini et al. 1999), labrids (Heiser et al.
2000), scarids (Moura et al. 2001, Gasparini et al. 2003), dactyloscopids (Feitoza 2002), labrisomids (Sazima
et al. 2002, Guimarães & Bacellar 2002), gobiids (Sazima et al. 1997), microdesmids (Gasparini et al. 2001),
and tetraodontids (Moura & Castro 2002). Several valuable species of Caribbean fishes that were long
assumed to extend to Brazil have also been shown to not occur there. Examples include Nassau grouper (Epi-
nephelus striatus), hogfish (Lachnolaimus maximus), and several parrotfish species (Sadovy & Eklund 1999,
Moura et al. 2001, Moura & Sazima 2003).
Endemism levels in the tropical southwestern Atlantic are therefore higher than previously supposed
(Briggs 1974), encompassing a broad taxonomic spectrum of pelagic (e.g. Menezes 1971), estuarine (e.g.
Bemvenuti 1995), demersal (e.g. Menezes & Figueiredo 1998), and reef-associated fishes (e.g. Moura & Saz-
Counts and measurements follow Anderson (1967) and Allen (1985); SL refers to standard length and HL to
head length. Measurements were made with dial calipers and recorded to the nearest 0.1 mm. Type specimens
are in the fish collections of MZUSP and USNM (institutional abbreviations follow Leviton et al. 1985). The
number of specimens, followed by the size range in mm SL, is given in parentheses following each catalog
number. Description is based on the holotype and all 21 paratypes. In the description, ranges of counts and
measurements are presented first, followed by values for the holotype in parentheses. Proportional measure-
ments are either expressed as percentages of standard length (% SL) or head length (% HL). Specimens were
collected on reefs with SCUBA using nets and speargun or with hook and line. Underwater photographs were
obtained to document live color patterns. In shallow patch reefs, tidepools, mangrove roots and grassbeds,
seine and hand nets were used for collecting. In addition to the type material described below, we examined
specimens of all Lutjanus species known from the southwestern Atlantic (see Comparative Material section).
Vertebral count was obtained from a radiograph of the holotype.
Lutjanus analis: MZUSP 66038 (1, 90.2), São Sebastião (23°49’S, 45°25’W), São Paulo State, Brazil. Lutja-
nus apodus: MZUSP 65929 (2, 142–138), Bahamas, San Salvador Island. Lutjanus bucanella: MZUSP 65240
(1, 232), Tamandaré (8°49’S, 35°5’W), Pernambuco State, Brazil. Lutjanus cyanopterus: MZUSP 65931 (1,
Lutjanus apodus not of Walbaum, 1792. Jordan, 1891:330 (Bahia); Ribeiro, 1915 (not paginated); Jordan & Fessler,
1893:510 (Pernambuco); Gilbert, 1900:170; Starks, 1913:32; Fowler, 1941:160; Allen, 1985:56 (in part); Carvalho-
Filho, 1994:138 (in part); Rocha et al., 1998:562; Rocha & Rosa, 2001:990; Moura & Menezes, 2003:82 (in part).
Lutjanus griseus not of Linnaeus, 1758. Ribeiro, 1915 (not paginated, in part); Ribeiro, 1918 (not paginated, in part);
Menezes & Figueiredo, 1980:22 (in part), Carvalho-Filho, 1994:135 (in part); Moura & Menezes, 2003:83 (in part).
Holotype: MZUSP 65664 (153) Camurupim Reef, Tamandaré (8°49'S, 35°5'W), State of Pernambuco, Brazil,
05 m depth; collected by R.L. Moura, R.B. Francini-Filho & C.H. Flesh, 22 January 1999.
Paratypes (21 specimens, 69–243): USNM 388233 (1, 161), Abrolhos Bank (17°57’08”S, 38°41’W),
State of Bahia, Brazil, 20 m depth, collected by R.L. Moura & R.B. Francini-Filho, 1 April 2000. MZUSP
52721 (2, 227–243) Parcel dos Abrolhos (17°57’08 S”, 38°41’6”W), Abrolhos Bank, State of Bahia, Brazil,
collected by R.L. Moura, R.B. Francini-Filho, C. Sazima & I. Sazima, 22 January 1998. MZUSP 84862 (1,
108), collected with the holotype. MZUSP 66066 (1, 131) Itaparica (12°52’S, 38°41’W), State of Bahia, Bra-
zil, collected by A. Carvalho-Filho, October 1982. MZUSP 60838 (1, 150) Parcel das Paredes (17°53’54”S,
38°57’13”W), Abrolhos Bank, State of Bahia, Brazil, collected by R.L. Moura, C.E.L. Ferreira & R.B. Fran-
cini-Filho, 19 February 2000. MZUSP 66021 (3, 82–95) Salvador (12°56’S, 38°31’W), State of Bahia, Brazil,
collected by V.G. Almeida, 21 July 1970. MZUSP 65939 (3, 71–89) same locality as MZUSP 66021, collected
by C.E. Dawson, N.A. Menezes & V.G. Almeida. MZUSP 66020 (1, 82) same locality as MZUSP 66066, col-
lected by N.A. Menezes & V.G. Almeida, 24 August 1972. MZUSP 66022 (5, 69–128), Lagoa Mundaú
(09°37’S, 35°48'W), Maceió, State of Alagoas, Brazil, collected by CETESB, 1985. MZUSP 66026 (1, 79)
Ponta de Pedras (8°3’S, 34°46’W) State of Pernambuco, Brazil, collected by P. Montouchet, 30 August 1970.
MZUSP 51167 (1, 87), same data as MZUSP 66022. MZUSP 65062 (1, 91) Fortaleza (3°45'S 38°20'W) State
of Ceará, Brazil, 0.5m depth (tidepool), collected by R.L. Moura, R.B. Francini-Filho & C.H. Flesh, 26 March
2000.
Diagnosis. Among western Atlantic snappers, Lutjanus alexandrei is similar to the gray snapper, Lutjanus
griseus (Linnaeus), and the schoolmaster, Lutjanus apodus (Walbaum), with which it shares the following
characters: 14 soft dorsal-fin rays, 6 (5–7 in L. apodus) scales between lateral line and dorsal-fin origin,
anchor-shaped vomerine tooth patch with a median posterior extension, upper canines much larger than lower
canines. Lutjanus alexandrei differs from L. griseus by the presence of 6 pale vertical bars (sometimes faded)
on the trunk (absent in L. griseus), a longer pectoral fin (length exceeding the distance from the snout to the
posterior margin of preopercle), and the angularity of the dorsal scale rows below the soft dorsal fin (not
markedly oblique as in L. griseus). Lutjanus alexandrei differs from L. apodus in having a red to pale-red
body cast and reddish fins instead of a yellowish-pale body with yellow fins, and by having only six narrow
pale vertical lines dorsally (sometimes faded) instead of eight wider pale bands as in L. apodus. The number
of lateral-line scales, varying from 43 to 48, is also slightly higher than that of L. apodus (40–45), and more
similar to that of L. griseus (43–47).
Description. The general morphological features of Lutjanus are presented by Allen (1985) and Anderson
(1987) for adults and by Lindeman et al. (2005) for larvae and juveniles, and are not repeated herein. Dorsal
rays X, 14; anal rays III, 8; pectoral rays 16 (uppermost rudimentary and unbranched); pelvic rays I, 5; princi-
pal caudal rays 9+8; lateral line with 43–48 (45) pored or tubed scales forming a continuous sensory canal
between the upper end of gill opening and the caudal-fin base, tubes in lateral line simple, unbranched; bran-
chiostegal rays 7; gill rakers 17–19 (17) total, 5–6 (5) fully developed rakers on the upper limb and 6–9 (8)
fully developed plus 3–5 (4) rudimentary rakers on the lower limb; gill membranes separate, free from isth-
TABLE 1. Proportional measurements for the holotype and 21 paratypes of Lutjanus alexandrei, sp.n. Measurements 1
to 6 expressed as proportions of standard length, and measurements 7 to 22 expressed as proportions of head length.
TABLE 2. Frequency of counts for some type specimens of Lutjanus alexandrei, sp. n. (specimens with damaged gill
rakers/ lateral-line scales not included).
Origin of dorsal fin slightly posterior to opercular margin; dorsal fin continuous, with only a slight notch
between the spinous and soft portions. Caudal fin slightly emarginated; anal fin rounded; pectoral fin pointed
and reaching the level of anus, length longer than the distance from tip of snout to posterior edge of preoper-
cle, and 3.0–3.6 (3.5) times in SL. Nostrils small, posterior nostril elongate and at level of pupil, anterior nos-
tril more rounded and located slightly below the level of the posterior one. Preopercle serrate, its lower margin
FIGURE 1. Holotype of Lutjanus alexandrei, MZUSP 65664, 152.6 mm SL, Camurupim Reef, Tamandaré (8°49'S,
35°05'W), State of Pernambuco, Brazil.
FIGURE 3. Early juvenile individual of Lutjanus alexandrei, 27 mm SL, collected in the mouth of Rio Mamucabas,
Tamandaré (08°49'S, 035°05'W), State of Pernambuco, Brazil, 1 m depth (Beatrice P. Ferreira & Sérgio Resende, 18 Feb-
ruary 2005).
Distribution, ecology and behavior. The Brazilian snapper, L. alexandrei is only recorded from the trop-
ical portion of the southwestern Atlantic continental shelf, and has a narrower latitudinal range than other
Western Atlantic species of Lutjanus. It is known from the state of Maranhão (00°52’S) to the southern coast
of the state of Bahia (18°0’S), Brazil, in areas under the influence of the west-flowing Equatorial Current
(northern Brazil) and the south-flowing Brazil Current (northeastern Brazil). It is apparently absent from oce-
anic islands. Additional collections may show an even broader distributional range for this species, as was the
case with 48 other poorly known reef-fish species in the southwestern Atlantic (Moura et al. 1999).
Habitats of the Brazilian snapper include coral reefs, rocky shores, coastal lagoons with brackish water,
mangroves and other shallow habitats with a mixture of soft- and hard-bottom. Recorded depths range from
intertidal (early stages only) to at least 54 m (Feitoza et al. 2005 — identified as L. apodus). During the day,
adults of L. alexandrei were observed on reefs as solitary individuals or in small groups showing restricted
activity. Adults can co-occur with L. jocu (see figure on page 40 in MMA 2002, several L. alexandrei were
misidentified as L. jocu). These mixed groups are often composed of large (> 20 cm), probably adult, individ-
uals. Similar to several other Lutjanus species, this species appears to be active predominantly during crepus-
cular and nocturnal periods. Juveniles smaller than 10 cm SL can be common in mangroves and rocky
Discussion
The existing literature typically gives the southern distributional limits of L. griseus and L. apodus as south-
eastern Brazil and northeastern Brazil, respectively (e.g. Menezes & Figueiredo 1980, Uyeno et al. 1983,
Allen 1985, Moura & Menezes 2003). All examined lots labeled as Lutjanus griseus and L. apodus in the two
major marine fish collections in Brazil (MZUSP and MNRJ) are actually L. alexandrei. We have determined
from photographs or voucher materials that all lutjanids commonly identified as L. griseus or L. apodus in
field surveys also are L. alexandrei. The same result applies for reef and estuarine fishery catches. The pres-
ence of either species in Brazil is highly doubtful given their absence in Brazilian museum collections, field
surveys, and fishery landings.
In the southwestern Atlantic there are cases of Caribbean fishes restricted to offshore islands [e.g. Inermia
vitatta (Inermiidae) and Haemulon chrysargyreum (Haemulidae)] (Moura & Sazima 2003) or occurring only
along restricted portions of the coast [e.g. Chromis scotti (Pomacentridae)] (Moura et al. 1999, Rocha & Rosa
2001). With more surveys, small populations of L. griseus or L. apodus could be definitively recorded from
offshore or mainland sites in Brazil. Clearly, the abundant and widespread L. alexandrei is not a product of
current hybridization of L. griseus and L. apodus, which, if present in Brazil, must be highly uncommon.
Other prominent reef species have also been erroneously recorded from Brazil, including Epinephelus striatus
and Lachnolaimus maximus (Serranidae and Labridae) (Sadovy and Eklund 1999, Moura & Sazima 2003).
Careful examination of a wide array of seemingly pan-western Atlantic reef species may result in additional
changes to long-assumed southern distributions.
Recent discoveries of new species belonging to major coastal fish families in Brazil (referenced in Intro-
duction) suggest a considerable level of isolation between Caribbean and Brazilian populations of coastal
fishes, especially those that associate with reefs. Genetic studies also indicate limited gene flow between these
two zoogeographic areas, even within groups with a high potential for dispersion such as surgeonfishes
(Rocha et al. 2002). Mechanisms of speciation and faunal enrichment in another group of coastal fishes (Scar-
idae) were recently studied in the Atlantic by Robertson et al (2006), based on mtDNA and nDNA sequences
to assess the effects of two major geographic barriers (the >30 million year old Atlantic ocean and the ~11
million year old Amazon–Orinoco outflow). Fluctuations in sea level, climate, and ocean-current dynamics
over the past ~10 million years likely produced marked variation in the effectiveness of the Amazon barrier,
allowing intermittent dispersal leading to establishment and allopatric speciation. This dynamic Amazon bar-
rier represents a major engine of West Atlantic faunal enrichment that has repeatedly facilitated bi-directional
dispersal, allopatric speciation, and remixing of the Caribbean and Brazilian faunas (Moura & Sazima 2003,
Rocha 2004, Robertson et al. 2006).
Because of this considerable isolation, caution should be used when management decisions are based on
populations from different sides of the Amazon-Orinoco barrier. Moreover, several of the recently discovered
species are relatively abundant on coastal habitats such as reefs and mangroves, but were frequently misiden-
tified (see Rocha & Rosa 1999, Moura et al. 2001). This amplifies the need for depositing voucher specimens
from ecological and fishery studies in major collections, supports the call for collection building in developing
countries, and stresses the need for additional analyses of the numerous systematic problems peculiar to tropi-
cal coastal ichthyofaunas.
Bodianus aya Bloch, 1790, described from Brazil after a pre-Linnaean work by the Dutch naturalist Marc-
Key to the Western Atlantic species of Lutjanus (modified from Allen 1985 and Anderson 2003)
1a. Dorsal fin with 10 spines and usually12 soft rays (rarely 11 or 13) ............................................................ 2
1b. Dorsal fin usually with 10 spines (rarely 9 or 11) and 14 soft rays (rarely 13 or 15)................................. 3
2a. Gill rakers 7–8 + 15–17 (including rudiments); 1/4 to 1/2 of black lateral spot extending below lateral
line; no evident paler (yellow in life) stripes on body and head; angle of preopercle greatly projecting and
strongly serrated...........................................................................................................L. mahogoni (Cuvier)
[southeastern coast of Florida and eastern coast of Yucatan peninsula to Venezuela]
2b. Gill rakers 6–7 + 13–14 (including rudiments); less than 1/4 of black lateral spot extending below lateral
line; 8–10 prominent paler (golden-yellow in life) stripes on body, often also 3–4 irregular paler (golden-
yellow in life) stripes on head; angle of preopercle scarcely projecting and finely serrated.........................
................................................................................................................................... L. synagris (Linnaeus)
[Bermuda and North Carolina to Santa Catarina, southern Brazil, including the West Indies, Gulf of Mex-
ico, and Caribbean Sea]
3a. Large, pronounced black spot at base and in axil of pectoral fin in specimens larger than 5 cm SL; dark
area at base of soft portion of dorsal fin (not always visible on preserved specimens); no black spot on
flanks; anal fin rounded ............................................................................................ L. buccanella (Cuvier)
Acknowledgments
We are indebted to William D. Anderson Jr. (GMBL) for providing us with literature and data, and for review-
ing the manuscript. Ronaldo B. Francini-Filho (Universidade Federal da Bahia), Naércio A. Menezes and José
Lima de Figueiredo (MZUSP), Rodolfo Claro (Instituto de Oceanología, Cuba), Ricardo M. C. Castro (Labo-
ratório de Ictiologia, Universidade de São Paulo, Ribeirão Preto), Biliana Cicin-Sain (University of Dela-
ware), Les Kaufman (Boston University), Gerald R. Allen and Guilherme F. Dutra (Conservation
International), Matheus O. Freitas and Maurício Hostim (VIDAMAR, Projeto Meros), Beatrice P. Ferreira and
Sérgio Resende (Universidade Federal de Pernambuco), Carolina V. Minte-Vera (UEM/NUPELIA), Jeff
Clayton (USNM) and Luiz A. Rocha (University of Hawaii) provided insightful comments, unpublished
information, and/or logistical support. Mário de Pinna (MZUSP) provided the radiographs and Paulo A.
Buckup allowed us to examine material deposited at MNRJ. This paper benefited from a PRONEX/MCT
grant to Naércio A. Menezes, a FAPESP grant (98/03491-3) to RLM, travel funds from CI-Brasil and ED to
RLM and KCL, travel funds from the U.S.-Brazil Marine Policy Consortium via the U.S. Department of Edu-
cation and the Coordenadoria de Aperfeicoamento de Pessoal de Nivel Superior (CAPES) to KCL. This is
Contribution Number One of the Marine Managed Areas Science Program, Brazil Node.
Literature cited
Allen, G.R. (1985) FAO species catalogue. Vol. 6 Snappers of the world: An annotated and illustrated catalogue of the
lutjanid species known to date. FAO Fisheries Synopsis 125, 1–208.
Allen, G.R. (1987) Synopsis of the circumtropical genus Lutjanus (Pisces: Lutjanidae). In: Polovina, J.J. & Ralson, S.