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Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5249.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:A19BA431-94FE-47A1-9C35-E4BB426ED4DE
Abstract
One new species of Neotropical Premnobius Eichhoff is described: P. brownei Atkinson & Flechtmann from Brazil. The
presence of the related genus Premnophilus Browne is confirmed from South America and six new species are described:
P. bertii Atkinson & Flechtmann from Brazil, P. jordali Petrov from Peru, P. maiai Atkinson & Flechtmann from Brazil,
P. pedrosai Atkinson & Flechtmann from Brazil, P. sarahsmithae Atkinson & Flechtmann from Brazil and Trinidad and
Tobago, and P. wilsoni Atkinson & Flechtmann from Brazil. French Guiana, and Peru. Gnathotrupes megapunctatus
Bright, 2019 is recognized as a synonym of Premnobius perezdelacruceii Petrov & Atkinson, 2018. A synopsis of
previously described Premnobius species and keys to species in both genera are presented. Generic characters for the
three genera of Neotropical Ipini are discussed.
Key words: Premnobius, Premnophilus, taxonomy, Acanthotomicus, generic characters, South America
Introduction
Previous work by Petrov & Atkinson (2018) and Atkinson et al. (2018) confirmed the existence of four native
Neotropical species of the genus Premnobius Eichhoff, previously considered to be exclusively Afrotropical, with
two species introduced into the Americas. Cognato (2013) moved Premnobius from the Xyleborini to the Ipini based
on molecular and morphological evidence. He also resurrected the genus Premnophilus Browne from synonymy
for two African species. In the same paper he also noted the presence of an undescribed species of Premnophilus
from Guyana. Cognato (2013) placed both genera in a separate subtribe within the Ipini, the Premnobiina Browne,
following the original suggestion by Browne (1962).
Review of specimens from Brazil and Peru revealed the existence of new species within the Premnobiina,
notably species not assignable to Premnobius or Acanthotomicus Blandford. Characters given by Browne (1962)
and Cognato (2013) allowed us to identify them as members of Premnophilus.
Existing keys to genera (Wood 1982, 1986, 2007) treat Premnobius (including Premnophilus) in the Xyleborini.
Even to an untrained observer, specimens of Premnobius and Premnophilus do not resemble any of the Neotropical
genera of Xyleborini. Their similarity to other Ipini, especially to Neotropical species of Acanthotomicus, is apparent.
We discuss here diagnoses for these genera in the Americas.
Because of recent changes in the tribal placement of the Neotropical Premnobiina, new synonymy, and
new species, we include a synopsis of all Neotropical species. Because all species are illustrated here only short
descriptions are included for previously described species. Our understanding of the Neotropical Premnobiina is very
preliminary. The Premnophilus species described here are known from a limited number of specimens collected in
traps with no knowledge of hosts or biology. There is no published phylogenetic information on Neotropical species
although Cognato (2013) found that an unnamed species of Premnophilus was nested within the African species.
All specimens cited are deposited in the collections listed below. All examined specimens were female.
Photographs were taken with a Canon camera with a Cannon MP–E 65 mm 1–5X macro lens mounted on a
Cognisys Stackshot rail system. Images were stacked with Zerene Stacker.
We are following a strictly morphological species concept here because there is no additional genetic, behavioral
or ecological information available on the species treated. Previously we were aware of several of the Premnophilus
species described here but were uncertain of their generic placement. Originally we had entertained the idea that
they might belong in the Xyleborini or Corthylini: Pityophthorina. A natural concern is whether any of them have
previously been described in other genera. The first author has examined types or authoritatively determined
specimens in the Wood (USNM) and Schedl (NHMW) collections of all taxa reported from the Neotropics for the
Xyleborini, Dryocoetini, and Trypophloeini. There are 213 species of Pityophthorina that have been described from
South America, mostly in the genera Pityophthorus Eichhoff and Araptus Eichhoff. More than 3/4 of these have
been photographed or can be discarded from descriptions. A revision of the Neotropical Pityophthorina would be an
enormous undertaking. In summary, we believe that the probabilities of previous description are low.
Taxonomy
Diagnostic characters of the three ipine genera found in the American tropics are shown in Table 1. This summary
is based on species found in the Americas, native or otherwise, and may not reflect characters of species in these
genera found on other continents. Species of Premnobius and Premnophilus are found in both the Neotropical and
Afrotropical regions. Acanthotomicus, as treated by Wood (1986), includes species from those regions as well as
from tropical and subtropical Asia.
Figure 1. Antennae of genera of Neotropical Premnobiina. A. Premnobius brownei Atkinson & Flechtmann; B. Premnophilus
maiai Atkinson & Flechtmann; C. Acanthotomicus granulatus (Ferrari). Photos by T.H. Atkinson (A,B) and Sarah Smith (C,
property of the Smithsonian Institution, used with permission).
These authors also stated that the club of Premnophilus is obliquely truncate while flattened in Premnophlius.
This latter character is difficult to interpret and seems variable to us. The term “obliquely truncated as widely used
in classification of the Scolytinae refers in part to the fact that in many groups the sutures on the anterior face of
the club are displaced distally on the posterior face. In many genera of the Ipini, Dryocoetini and Xyleborini the
basal part of the antennal club (proximal to the first suture) is sclerotized or corneus with the remainder of the club
mostly pubescent. In specimens where it is possible to see the antennal club laterally, this displacement is apparent.
It seems to us that this is more of a relative than an absolute distinction. In American species of Premnophilus the
basal corneous portion on the posterior face of the club covers 3/4 of the face and any pubescent area is distal to it.
In both Premnobius and Acanthotomicus the corneous portion occupies less than half of the length of the club and
other sutures are visible on the lateral areas, nearly to the base.
There are also notable differences in the declivity of American species. Cognato (2013) pointed out that the
posterolateral margin of the declivity is rounded in Premnophilus and acutely margined in Premnobius, and this
is true of American species of Premnophilus (Figs. 6–12). In some American species of Premnobius the declivital
margin is not as acute basally (Figs. 4–5), but it is well marked and sometimes granulate. In most American species
of Acanthotomicus the declivital margin is very well defined and elevated (Fig. 2). The shape of the declivity is also
useful in distinguishing these genera, specifically the steepness as seen in lateral profile and degree of concavity
on the declivital face. In Acanthotomicus the declivity is very steep and the face is strongly or weakly concave. In
Premnophlius the declivity is steep with the face flattened or slightly sulcate. In species of Premnobius the declivity
is longer, proportional to the elytral length and generally concave or flattened.
The ratio of the total body length to that of the maximum body width is commonly used in species level
identification. Ranges are taken from the present study and published sources (Wood 1982, 1985, 2007; Petrov &
Atkinson 2018; Atkinson et al. 2018). Species of Premnobius are much more slender than those of the other two
genera.
The following key will distinguish females of all species of Premnobius known from the Americas, both native and
exotic.
1 Declivity with a pair of quadrate spines on lower lateral margins, their height subequal to width at base; spines on lateral
margins often blunt or digitate (Fig. 4 A–H) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Declivity without pair of quadrate spines on lower margins; all granules or spines on lateral margins acutely pointed or digitate
(Fig. 5 A–F) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
2 Pronotum not elongate, length 1/4 of body length; quadrate elevation near middle of declivity in lateral view (Fig. 4 A,B,E,F)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Pronotum elongate, length 1/3 of body length; quadrate elevations on posterior 1/3 of declivity in lateral view (Figs. 4 C,D,
G–H). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3 Quadrate elevation thickened basally, displaced medially from posterolateral crest; prominent, acute granules at base of declivity
on interstriae 2 and 4 (Fig. 4 A,B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . flechtmanni (Wood)
- Quadrate elevation not thickened, on posterolateral crest, granules at base of declivity inconspicuous (Fig. 4 E,F). . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . neoadjunctus (Schedl)
4 Acute denticle near base of declivity on interstria 3 short, displaced medially from declivital crest; length 3 mm (Fig. 4 C,D). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . perezdelacrucei Petrov & Atkinson
- Acute denticle near base of declivity on interstria 3 digitate, nearly as long as quadrate process, on declivital crest ; length 2 mm
Fig. 4 G,H). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . assiduus (Schedl)
5 Large digitate spine near apex of declivity, its length at least 3 its basal diameter (Fig. 5 A–B). . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brownei Atkinson & Flechtmann
- Declivital margin without large spines near apex, either without large spines or largest spines near midpoint of declivity (Fig. 5
A–D). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6 Interstria 1 on declivity with a row of small pointed tubercles; raised lateral margin of declivity with small pointed granules;
transition from elytral disc to declivity pronounced, not gradual (Fig. 5 A,B) . . . . . . . . . . . . . . . . . . . . . . . . cavipennis Eichhoff
- Interstria 1 on declivity without any granules; several large, pointed tubercles near base and midpoint of declivity on lateral
margin; base of declivity extends gradually anteriorad at base along insterstria 1 (Fig. 5, C,D). . . . . . . . ambitiosus (Schaufuss)
Similar species. This is the largest and most robust species found in the Neotropical region. It most closely resembles
P. cavipennis, but is much larger.
Similar species. This species most closely resembles P. perezdelacrucei in that both are very elongate with
particularly long pronota.
Diagnosis. Length 1.80–2.25 mm, 3.6–4.4 × as long as wide (Atkinson et al. 2018: 2). Frons with small,
sparse punctures, vestiture more abundant laterally and on epistoma. Pronotum elongate, almost 2 × as long as
wide; summit well before middle, anterior slope steep, disc about 75% of pronotal length, shining, punctures fine,
vestiture sparse. Strial punctures on elytral disc not impressed, interstriae about 2 × as wide as striae. Interstrial
setae fine, uniseriate, length slightly longer than interstrial width, becoming longer towards declivity. Declivity
occupying 33% length of elytra, shallowly concave, with an elevated crest from interstria 4 to apex, transition
gradual anteriorly. Armature consisting of digitate projection on crest at interstria 4, its length 3 × its width at base;
a larger quadrate, blunt projection, laterally compressed, slightly displaced mesad from crest on lower fourth; small,
irregular granules also present. Punctures confused on declivital face, deeper than those on disc, some short, slender
setae, these shorter than those near declivital crest.
Distribution. Northern South America and Amazon Basin (Fig. 12D).
New Records. Brazil: Amazonas, Manaus, Universidade Federal do Amazonas, Campus Senador Arthur
Virgilio Filho, 35.57′ S, 5957.674′W, 24-VI-2021, ethanol-baited FIT, E.A.C. Corrêa, dense ombrophilus forest
(MEFEIS, 1); Venezuela, Mérida, 20 km SW El Vigía, 50 m, 10-XII-1989, tree seedling, collection 180, S.L. Wood
(USNM, 1, in unidentified material in Wood collection).
Diagnosis: Among the American species P. brownei is unique in that the large process on the declivital crest is
elongate and digitate, rather than quadrate..
Female. Total length: 1.13 mm, maximum width: 0.35, length of elytra: 0.65 mm, total length / width: 3.23,
elytra length / width: 1.86, pronotal length / width: 1.37 (n=1). Color dark reddish brown, darker on posterior half
and declivity.
Frons smooth, shining, surface shallowly punctured, punctures separated by 3 their diameters, a weak
longitudinal carina evident from above the upper level of the eyes to the epistoma. First segment of the anterior face
Figure 3. Premnobius brownei Atkinson & Flechtmann, female holotype. A. dorsal habitus; B. lateral habitus; C. frons and
antenna; D. declivity. All photographs by T.H. Atkinson.
Similar species. Among the species found in the Americas, this most closely resembles P. ambitiosus.
Diagnosis. Length 2.3–2.8 mm, 3.3 as long as wide (Wood 2007: 368). Frons granulate, punctate, vestiture
sparse. Anterior margin of pronotum rounded, unarmed. Pronotum about half as long as elytra, summit slightly
anterior to middle, anterior asperities short, abundant. Disc shining, with shallow widely spaced puncture; short,
semi-recumbent setae. Elytral striae not impressed; interstriae 3 as wide as striae; interstrial setae confused to base,
length shorter than interstrial width, slightly more abundant near declivity. Declivity occupying posterior 25% of
elytral with a nearly complete circumdeclivital elevated crest, granulate along crest, none notably larger than others.
Face of declivity concave, strial and interstrial punctures much larger than those on disc, confused, some along
sutural interstriae granulate; some setae present, notably along suture.
Distribution. Introduced from the Afrotropical region. It is widely distributed in most tropical and subtropical
regions of the Americas (Fig. 12A). It is commonly collected in light traps and flight intercept traps baited with
ethanol.
Similar species. This species closely resembles P. neoadjunctus. At present very few specimens of either species
have been collected. Further collection is needed to determine whether these are distinct species.
Diagnosis. Length 2.0 mm, 3.0 as long as wide (Wood 2007: 337). Frons reticulate in center, punctate laterally.
Anterior margin of pronotum rounded, with 8–10 small teeth along margin. Pronotal summit about 33% of pronotal
length. Disc shining, with shallow, widely spaced punctures and very short, erect setae, their length less than
distance between punctures. Striae on disc not impressed, interstriae 3 as wide as striae. Interstrial setae uniseriate,
their length slightly longer than interstrial width, longer and stouter on odd-numbered interstriae, becoming longer
posteriorly. Declivity occupying 33% of elytral length, shallowly concave. Raised, acute declivital crest extending
from interstria 4 to apex. A small, acute denticle present on interstria 2 at base of declivity; a larger one on interstria
4, displaced mesally from crest, and a quadrate, blunt, flattened projection near middle of declivity, also displaced
Similar species. This species most resembles P. flechtmanni in that both have only a single blunt projection on the
declivital crest without additional tubercles.
Diagnosis. Length 2.8 mm, 3 as long as wide (Wood 2007: 670). Frons granulate-punctate. Anterior margin of
pronotum subtruncate, unarmed. Pronotum with summit at anterior third, disc shining with moderately deep punctures,
separated by 3–4 their diameters, vestiture of short, fine, erect setae, longer laterally. Striae on disc not impressed;
interstriae 3 as wide of striae. Uniseriate interstrial setae fine, about as long as width of interstriae, becoming much
longer posteriorly near declivital crest. Declivity occupying posterior third of elytra, concave, with an elevated crest
from interstria 4 to apex, transition gradual at base. Declivital crest irregularly granulate, with a quadrate, blunt, laterally
compressed process about 2/3 distance from base to apex. Strial punctures distinct, uniseriate on face of declivity, large
than on disc. Long setae on crest also present in anterolateral areas of face, before quadrate processes.
Note. Atkinson et al. (2018) mistakenly showed the declivity of P. assiduus in Fig. 4D, not the declivity of this species.
Distribution. Known only from two localities in Amazonian and southern Brazil (Fig. 12D).
Similar species. Among the American species this most closely resembles P. assiduus.
Diagnosis. Length 2.45 mm, 4 as long as wide (Petrov & Atkinson 2018: 42). Very slender, with proportionately
elongate pronotum. Frons shining, weakly reticulate, with sparse setae. Pronotum elongate, 80% as long as elytra;
anterior margin rounded, weakly elevated with small asperities; summit located about 25% of length from anterior
margin. Disc smooth, shining, with shallow, sparse punctures; vestiture a mixture of taller and shorter erect setae.
Elytral striae not impressed, interstriae 2 width of striae; vestiture of erect setae, length greater than width of
interstriae, mostly uniseriate, about twice as long at base of declivity. Declivity occupying posterior 33% of elytral
length. A raised posterolateral crest present from interstria 4 to apex, transition gradual above. Declivity with a
small, sharp tooth near base on interstria 3, displaced mesally from declivital margin; a laterally flattened, apically
rounded projection on crest about 2/3 of the distance to apex, its basal length greater than its height.
New Records. Brazil: Bahia, Una, Fazenda Vera Cruz, 1518′24″ S 3909′30″W, 24-II-2013, ethanol-baited,
cocoa cabruca agroforestry system, S. Novais (MEFEIS, 1); Minas Gerais, Capinópolis, Fazenda Grama,
1846′7.32″S 4928′38.65″W, 16-III-2020, 30-III-2020, ethanol-baited FIT, Khaya grandifoliola stand planted IX-
2015, C.F. Faria (MEFEIS, 2); same general locality, 1846′6.18″S 4928′51.59″W, 10-III-2020, ethanol-baited,
Khaya grandifoliola stand planted IX-2014, C.F. Faria (MEFEIS, 1); French Guiana: Paracou (Res. Station?), XI-
1996, canopy window trap, P.G. Hammond (NHMUK, 1).
Distribution. Widely distributed from southeastern Mexico and the Dominican Republic to southern Brazil (Fig. 12C).
Type species. Xyleborus joveri Schedl, 1951: 41 [=Xyleborus quadrispinosus Schedl, 1938: 461; synonymy Schedl
1963: 558]; original designation.
Taxonomic comments. Schedl (1964: 52) treated Premnophlius as a synonym of Premnobius. Cognato (2013)
removed Premnophilus from synonymy with Premnobius. He included the two Afrotropical species originally
placed there by Browne and also indicated that there was an unnamed species from Guyana.
Distribution. At present, two species are known from the Afrotropical region and seven new Neotropical
species are described here. The American species are all known from tropical rain forest areas of the Amazon Basin
and northeastern Atlantic coast.
Biology. Browne (1962) noted that Premnophilus quadrispinosus (Schedl) has an ambrosial habit. All other
species, including the newly described Neotropical ones have been collected in traps or canopy fogging and nothing
is known of their habits, thought they are presumed to be ambrosia beetles.
1 Posterolateral margin of declivity not raised, declivital face flattened or sulcate, marginal denticles absent (Figs. 7 E, 9 E, 10 E,
12 E). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Posterolateral margin of declivity raised, rounded, declivital face concave, marginal denticles present (Figs. 6 E, 8 E). . . . . . . 5
2 Declivital face flattened, all denticles confused, uniform in size (Figs. 11 E)wilsoni Atkinson & Flechtmann
- Declivital face weakly sulcate at base, denticles on interstria 1 generally larger than others, uniseriate (Fig. 7 E, 9 E, 10 E). . . 3
3 Vestiture on elytral disc mostly absent, granules on interstria 1 of declivity not noticeably larger than others (Fig. 8). . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . jordali Petrov
- Vestiture on elytral disc abundant to base, granules on interstria 1 clearly larger than others. . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Declivity sulcate from base to apex, margin elevated in middle (lateral view), apex truncate, larger granules only on interstria
1 (Fig. 11). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . sarahsmithae Atkinson & Flechtmann
- Declivity weakly sulcate at base, margin not elevated in middle (lateral view), apex appearing acute, larger granules on interstria
1 as well as on other interstriae at apex (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pedrosai Atkinson & Flechtmann
5 Smaller (1.8 mm), declivity narrowly concave, with a small granule on the declivital margin near the base and a similar one near
the apex (Fig. 6). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bertii Atkinson & Flechtmann
- Larger (2.1 mm), declivity more broadly concave, with an acute denticle on the declivital margin near the base and two closely
spaced denticles near the apex (Fig. 8). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . maiai Atkinson & Flechtmann
Diagnosis. This species and P. maiai differ from those previously described here in that the declivity is clearly
sulcate, with a rounded margin from the declivital apex almost to its base which bears 2 or 3 denticles. There are no
granules on the declivital face.
Female. Total length: 1.81 mm, maximum width: 0.66, length of elytra: 0.99 mm, total length / width: 2.74,
elytra length / width: 1.50, pronotal length / width: 1.24 (n=1). Color reddish brown.
Frons reticulate, with small, shallow punctures, these widely spaced, some with erect setae slightly longer
than the distance between punctures. Punctures becoming granulate near epistoma. A weakly elevated, longitudinal
carina runs from well above the eyes to the epistomal margin. First segment of anterior face of antennal club
corneus, bisinuate, middle portion not reaching middle of club; distal part densely pubescent without visible sutures;
pubescent area not reaching base of antennal laterally. Posterior face with corneous portion occupying 3/4 club
length, pubescent distally, lateral areas not pubescent.
Anterior margin of pronotum rounded, asperities on its leading edge slightly larger than those on the rest of the
anterior slope. Summit slightly anterior to middle, asperities short, with about 3 their thickness, becoming wider
and confluent towards summit. Pronotal disc shining; punctures shallow, small, widely spaced. Erect setae on disc,
spaced by about their length.
Elytral disc shining, smooth. Striae not impressed, punctures shallow, with short recumbent setae. Interstriae
about 2 width of striae with mostly uniseriate erect setae, their length slightly longer than interstrial width. Interstriae
becoming weakly granulate near base of declivity. Declivity abrupt, steep occupying 20% of declivital length in
lateral view; steep, with face sulcate from the base to the apex. A rounded posterolateral marginal elevation is present
on the declivity from apex to near the base. Declivity narrowly concave in center of declivity between interstria 3,
narrowed posteriorly. Strial punctures and interstrial setae present on declivital face. Two small, acute denticles are
present on rounded declivital margin, one near base on interstria 2 and a larger one near the middle on interstria 3.
Male. Unknown.
Type Material. Holotype, female, Brazil: Amazonas: São Sebastião do Uatumã, Reserva Biológica do Uatumã, 4–II–
2010, 148′18″ S, 5915′5″ W, 18–II–2010, unbaited flight intercept trap, R.S.L. Abreu, ombrophilous dense forest (MEFEIS).
Distribution. Known only from the lower Amazon Basin in tropical rain forest (Fig. 12F).
Etymology. This species is named in honor of Evoneo Berti Filho, retired professor at the Escola Superior de
Agricultura “Luiz de Queiroz, Universidade de São Paulo, Piracicaba campus, São Paulo, Brazil who was the first
to do research on Scolytinae in reforested areas in the state of São Paulo.
Diagnosis. The new species is closely related to Premnophilus pedrosai and P. sarahsmithiae but can be distinguished
by the absence of large granules on interstria 1 and the less abundant vestiture on the elytral disc.
Female. Total length: 1.45–1.72 mm, maximum width: 0.50–0.67, length of elytra: 0.81–1.05 mm, total length / width:
2.56–2.86, elytra length / total length: 1.36–1.69, pronotal length / width: 1.1–1.3, (n=3). Color reddish brown to dark brown.
Frons reticulate, with large, shallow punctures, epistomal margin granulate. A weakly elevated, longitudinal
carina runs from well above the eyes to central part of frons. Vestiture of short sparse pale hairs, longer and abundant
in epistomal margin above mandibles. Gena glabrous. Eyes shallowly emarginate, 2.2 as long as wide. Antenna brown.
First segment of anterior face of antennal club corneus, middle portion reaching about middle length of club.
Pronotum reddish brown to dark brown in anterior part. Base and anterior margins of pronotum rounded, lateral
margins subparallel in 1/2 of pronotal length. Asperities on its leading edge small, same size those on the rest of the
Figure 7. Premnophilus jordali Petrov, female holotype. A. dorsal habitus; B. lateral habitus; D. declivity posterolateral view;
D. declivity, posterior view. All photographs by A.V. Petrov.
Figure 8. Premnophilus maiai Atkinson & Flechtmann, female holotype. A. dorsal habitus; B. lateral habitus; C. frons and
antenna; D. declivity. All photographs by T.H. Atkinson.
Diagnosis. This species and P. bertii differ from those previously described here in that the declivity is clearly
sulcate, with a rounded margin from the declivital apex almost to its base which bears 2–3 denticles. There are no
granules on the declivital face.
Female. Total length: 2.10 mm, maximum width: 0.75 mm, length of elytra: 1.18 mm, total length / width: 2.80,
elytra length / width: 1.57, pronotal length / width: 1.23 (n=1). Elytra reddish brown, pronotum yellowish brown.
Frons shining, smooth, with small, shallow punctures, these widely spaced, some with erect setae slightly
longer than the distance between punctures. Punctures becoming granulate near epistoma. A very weakly elevated,
longitudinal carina runs from well above the eyes but ends before the epistomal margin. First segment of anterior
face of antennal club corneus, bisinuate, middle portion not reaching middle of club; distal part densely pubescent
without visible sutures; pubescent area not reaching base of antennal laterally. Posterior face with corneous portion
occupying 3/4 club length, pubescent distally, lateral areas not pubescent.
Anterior margin of pronotum bluntly rounded, subtruncate, asperities on its leading edge notably larger than
those on the rest of the anterior slope. Summit slightly anterior to middle, asperities short, flattened, with about 3
their thickness, becoming shorter towards summit. Pronotal disc shining; punctures shallow, small, widely spaced.
Erect setae on disc, spaced by about their length.
Elytral disc shining, smooth. Striae not impressed, punctures shallow, with short recumbent setae. Interstriae
about 2 width of striae with mostly uniseriate erect setae, their length slightly longer than interstrial width. Interstriae
becoming weakly granulate near base of declivity. Declivity steep occupying 25% of declivital length in lateral view;
steep, with face sulcate from the base to the apex. A rounded posterolateral marginal elevation is present on the
declivity from apex to near the base. Declivity narrowly concave in center of declivity between interstria 3, narrowed
posteriorly. Strial punctures present on declivital face, surface dull. Three acute denticles are present on rounded
declivital margin, one near base on interstria 3 and a pair of closely spaced denticles closer to elytral apex.
Male. Unknown.
Type Material. Holotype, female, Brazil: Amapá, Tartarugalzinho, Comunidade Entre Rios—Projeto de
Assentamento Cedro, Retiro Paraíba, 17′32″ N, 5118′34″ W, 22–VIII–2015, FIT with ethanol, Amazonian terra
firme forest fragment, W.R. Silva (MEFEIS).
Distribution. Lower Amazonian region (Fig. 12F).
Etymology. This species is named in honor of Jos Luiz da Silva Maia, head of the Division of Forest Protection
and Environment of the forest company Duraflora in Agudos, state of São Paulo, where he worked for many years.
He worked collaboratively with the second author in Scolytinae research projects over many years.
Diagnosis. P. sarahsmithae, P. pedrosai and P. jordali are similar in that the declivity is flattened or slightly sulcate
without an elevated posterolateral crest. The declivity of all of these is covered with densely confused granules and
setae. This species differs from P. sarahsmithae in that the face of the declivity is more flattened, the larger granules
on interstria 1 are less regular and larger posteriorly. Similar granules are present in the apical regions of interstria 2.
Female. Total length: 1.42 mm, maximum width: 0.52 mm, length of elytra: 0.78 mm, total length / width: 2.73,
elytra length / width: 1.50, pronotal length / width: 1.23 (n=1). Color dark brown.
Frons reticulate, with large, shallow punctures, these contiguous, becoming granulate near epistomal margin.
A weakly elevated, shining, longitudinal carina runs from well above the eyes but does not reach epistomal margin.
First segment of anterior face of antennal club corneus, weakly bisinuate, middle portion reaching about 1/3 length
of club; distal part densely pubescent without visible sutures; pubescent area not reaching base of antennal laterally.
Posterior face with corneous portion occupying 3/4 club length, pubescent distally, lateral areas not pubescent.
Anterior margin of pronotum rounded, around ten asperities on its leading edge larger than those on the rest of
the anterior slope. Summit slightly anterior to middle, asperities short, with about 2 their thickness, becoming less
Figure 9. Premnophilus pedrosai Atkinson & Flechtmann, female holotype. A. dorsal habitus; B. lateral habitus; C. frons and
antenna; D. declivity. All photographs by T.H. Atkinson.
Figure 10. Premnophilus sarahsmithae Atkinson & Flechtmann, female holotype. A. dorsal habitus; B. lateral habitus; C.
frons and antenna; D. declivity. All photographs by T.H. Atkinson.
Diagnosis. P. sarahsmithae and P. pedrosai are similar in that the declivity is slightly sulcate without an elevated
posterolateral crest. This species differs from the other two because the declivity is weakly sulcate and the granules
are largest and in a uniseriate row on interstria 1.
Female. Total length: 1.45 mm, maximum width: 0.54 , length of elytra: 0.76 mm, total length / width: 2.69, elytra
length / total length: 1.41, pronotal length / width: 1.28 (n=1). Color yellowish brown, darker in declivital area.
Frons reticulate, with large, shallow punctures, these contiguous. A weakly elevated, shining, longitudinal
Diagnosis. This species differs from other species in that the declivity is completely flattened with a rounded,
circumdeclivital margin. Specimens from Peru differ slightly in the pattern of declivital vestiture.
Female. Total length: 1.31 mm, maximum width: 0.49, length of elytra: 0.69 mm, total length / width: 2.67,
elytra length / width: 1.41, pronotal length / width: 1.27 (n=1). Color yellowish brown.
Frons reticulate, with small, shallow punctures, these widely separated and with short, erect setae. A weakly
elevated, shining, longitudinal carina runs from well above the eyes to the middle of the frons, not reaching the
epistoma. First segment of anterior face of antennal club corneus, bisinuate, middle portion not reaching middle of
club; distal part densely pubescent without visible sutures; pubescent area not reaching base of antennal laterally.
Posterior face with corneous portion occupying 3/4 club length, pubescent distally, lateral areas not pubescent.
Anterior margin of pronotum rounded, asperities on its leading edge not larger than those on the rest of the
anterior slope. Summit slightly anterior to middle, asperities short, with about 3 their thickness, becoming wider and
confluent towards summit. Pronotal disc shining, faintly reticulate; punctures shallow, small, widely spaced. Sparse
erect setae on disc, spaced by about their length.
Elytral disc shining, smooth. Striae not impressed, punctures shallow, with short recumbent setae. Interstriae
about 2 width of striae with mostly uniseriate erect setae, their length slightly longer than the interstrial width. All
interstriae becoming granulate at the base of the declivity. Declivity occupying 25% of declivital length in lateral
Figure 11. Premnophilus wilsoni Atkinson & Flechtmann, female holotype. A. dorsal habitus; B. lateral habitus; C. frons and
antenna; D. declivity. All photographs by T.H. Atkinson.
We thank the Brazilian collectors who assisted in trapping surveys. We thank Anthony Cognato for his thorough and
knowledgeable comments. We thank Max Barclay (NHMUK) for the loan of material.
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