A new flea, Ectinorus (Ectinorus) insignis n. sp. (Siphonaptera, Rhopalopsyllidae, Parapsyllinae), with notes on the subgenus Ectinorus in Chile and comments on unciform sclerotization in the superfamily Malacopsylloidea.

A list is provided for the species of Ectinorus sensu stricto from Chile. Ectinorus (Ectinorus) insignis n. sp. is described from Chile: this species is characterized by the male genitalia. In the subgenus Ectinorus, the authors report the presence in Chile of E. pilosus Beaucournu & Carmen Castro, 2002 described from Argentina and E. simonsi (Rothschild, 1904) described from Bolivia but also known from Peru. A female neallotype is designated for E. ineptus Johnson, 1957. "Unciform sclerotization" is noted and illustrated for the first time, in all Malacopsylloidea, and a list is given for all studied species. © J.-C. Beaucournu et al., published by EDP Sciences, 2013.

Introduction

The interest of the genus Ectinorus (and the related genera Delostichus, Tetrapsyllus, etc.), regardless of its morphological features, is that it is one of the bubonic plague vectors, transmitting Yersinia pestis in Chile-Andean subregion (e.g., Macchiavello, 1948) [28]. Admittedly, the importance of plague in the southern cone of South America is of lesser importance than in the Old World (Kazakhstan, Mongolia, China, Vietnam, Republic of Congo, Tanzania, Madagascar), but the potential for disease remains and a knowledge of endemic fleas is important.

Genus Ectinorus Jordan, 1942

The genus Ectinorus (Rhopalopsyllidae: Parapsyllinae), endemic to the Chile-Andean subregion, includes related sigmodontine rodents fleas. Three subgenera are included in Smit’s review (1987) [36]: Panallius Jordan, 1942, Ectinorus sensu stricto and Ichyonus Smit, 1987. Until now 33 Ectinorus sensu stricto species were known (Hastriter & Sage, 2009) [16]. Fifteen species occur in Chile (those for which type localities are in Chile are marked with an asterisk) and include (listed alphabetically): E. chilensis* Lewis, 1976, E. cocyti* (Rothschild, 1904), E. curvatus* Beaucournu & Gallardo, 1991, E. gallardoi* Hastriter, 2001, E. ineptus Johnson, 1957 (in Beaucournu & Gallardo, 1991) [5], E. ixanus (Jordan, 1942) (in Beaucournu & Kelt, 1990) [9], E. lagidium* Beaucournu & Gonzàlez, 2005, E. levipes (Jordan & Rothschild, 1923) (in Beaucournu & Kelt, 1990), E. martini* Lewis, 1976, E. mimacydis* Beaucournu & Gallardo, 2004, E. mondacai* Hastriter, 2001, E. nomisis Smit, 1987 (in Beaucournu & Gallardo, 1989: description of female neallotype) [4], E. setosicornis Jordan, 1942 (in Beaucournu & Gallardo, 1991) [5], E. splendidus* Smit, 1968 and E. uncinatus* Beaucournu & Gallardo, 1991.

An additional three species are here reported from Chile for the first time: Ectinorus pilosus Beaucournu & Carmen Castro, 2002 [2] described from Argentina, E. simonsi (Rothschild, 1904) [32] known previously from Bolivia and Peru, and a new species, Ectinorus insignis Beaucournu & Gonzàlez-Acuña. In addition, a neallotype is designated for the female of Ectinorus ineptus Johnson, 1957 [19]. The total number of Ectinorus sensu stricto known from Chile is elevated to 18 species.

Ectinorus (Ectinorus) pilosus Beaucournu & Carmen Castro, 2002

Ectinorus pilosus Beaucournu & Carmen Castro, 2002: Description of holotype male and allotype female, Santa Maria (26° 40′ S–66° 02′ W) (Catamarca), Argentina, on Ctenomys sp. (later determined as Ctenomys knighti Thomas, 1919), collection date unknown.

For this species, one male and one female were found in Chile, Copiapó (Atacama), on Ctenomys sp., collection data unknown.

Ectinorus (Ectinorus) simonsi (Rothschild, 1904)

Pulex simonsi (Rothschild, 1904): two males and one female on Neoctodon simonsi (= Octodontomys gliroides) at Challapata (Avaroa, Bolivia), X/11/1901.

Rhopalopsyllus simonsi (Rothschild): Jordan & Rothschild, 1908 [23]: redescription.

Dysmicus acheronis Johnson, 1957 [19]: holotype male, Yuru (Arequipa, Peru) on Galea musteloides, VIII/8/1939. Smit (1987): acheronis synonymized with simonsi.

Ectinorus (Ectinorus) simonsi (Rothschild, 1904): drawings of male and female on Lagidium viscacia and a new locality is reported: Zudañez, Bolivia (in the Bolivian subregion, province of Jaime Zudañez), IV/19/1955 (Smit, 1987) [36].

One male found in Chile, Chusmiza (19° 40′ S–69° 10′ W), altitude 3430 m (Tarapaca), on Octodontomys gliroides, X/15/2011 (D. Gonzàlez-Acuña rec.). Identification is obvious for this species (apex of aedeagus, sternum VIII) but in this area on October 9, 1989, we found on the same host, at 2500 m altitude, three male intergrades between simonsi and nomisis Smit, 1987. These taxa are so close that it is likely that E. nomisis is a subspecies of E. simonsi.

Ectinorus (Ectinorus) ineptus Johnson, 1957

Ectinorus ineptus Johnson, 1957: description of male holotype (plates 67–69) at Picotani (Puno, Peru), on Phyllotis (Auliscomys) pictus, IX/14/1941.

Ectinorus (Ectinorus) ineptus Johnson 1957: Smit (1987) illustrated the genitalia of the male holotype (their Figures 189 and 190), from Auliscomys pictus.

Ectinorus (Ectinorus) ineptus Johnson, 1957, Beaucournu & Gallardo, 1991 [5]: Parinacota (18° 12′ S–69° 16′ W), altitude 4000 m, Chile, on Phyllotis darwini, IX/30/1989. One male and one female: Short description of the male and the female (Figures 6, 9, 12–14), female explicitly “not designated as neallotype because two other very close taxa have been collected in the same region syntopically with this species (e.g., E. curvatus n. sp.)”.

Ectinorus (Ectinorus) ineptus Johnson, 1957: Lake Chungara (18° 15′ S–69° 10′ W), altitude 4574 m, Chile (Parinacota) on Auliscomys boliviensis (Waterhouse, 1846), one male and one female XI/21/2011; Enquelga, altitude 3700 m, Chile (Tarapaca) on Eligmodontia puerulus (Philipi, 1896), one female X/17/2011 (D. Gonzàlez-Acuña rec.) (The determinations of all Chilean hosts are after Wilson & Reeder, 2005) [37].

So we have two pairs, each in syntopy on the same individual host and a sole female: therefore, the identity of the female drawn in 1991 is confirmed. Currently males, in Chile, have been collected from two locations close to each other.

We select the 1991 female, as neallotype, because the others have misplaced spermatheca, but are otherwise identical. So the neallotype is from Parinacota (18° 12′ S–69° 16′ W), altitude 4000 m, Chile (Parinacota) on Phyllotis darwini, IX/30/1989.

Description of females

Material examined: two males and three females, as indicated above.

Material deposited: Neallotype in the collection of first author (Collection JCB currently in Faculty of Medicine, Rennes), for ultimate deposition in Muséum National d’Histoire Naturelle, Paris, France (MNHN).

Head capsule: With well-developed tubercle, without micro-seta visible in the vallum. Genal margin slightly convex; maxillary palpus with segments I, II and IV of same length; segment III shorter; labial palpus with seven segments, not reaching apex of coxa; pre-antennal seta very small; six marginal setae and two small, thicker setae in posterior point of the gena; two occipital rows, respectively, of one and five setae.

Thorax: Prothorax with anterior row of five small setae and posterior row of seven with intercalaries. Mesothorax: 14 small setae and principal row of 6 longer setae; 10 pseudosetae. Metathorax: four anterior setae and posterior row of seven longer setae, most dorsal seta curved. Metepimeron with four long setae; spiracle pointed, or shaped as a “nightcap”. Femoral guard setae long; external half the length of internal. Tibia with eight notches on dorsal margin. Notches II, V and VIII with, at least, one long seta. Longest seta of notch VIII extends to apex of first tarsomere. Longest seta of this tarsomere extends to apex of tarsomere II; longest seta of tarsomere II extends to half of the length of distitarsomere.

Abdominal segments: Tergum I with eight spinelets, four on each side. Tergum II with row of seven long setae, most ventral seta at level of spiracle. Same setation on terga III to VII, but most ventral seta below spiracles; spiracles relatively large and rounded. Tergum VIII (Figure 2) with small area of seven or eight small setae anterior to spiracle; spiracle marginal and vermiform. Unciform sclerotization present (see below). Anal stylet (Figure 1) long, conical, with one small seta below insertion of long apical seta. Below ventral anal segment four long marginal setae and 16–18 lateral setae of various lengths and seven to eight thin, small mesal setae. Sternum II with 1 barely discernible striarium and laterally 17 thin lateral setae and 2 small marginal setae. Sternum III with lateral row of six setae and three to four dispersed setae. Sterna IV and V with main row of four or five setae and one isolated seta. Sternum VI with simple row of five setae. Sternum VII (Figure 2) with row of six setae inserted on half of the segment. Margin gently convex. Sternum VIII (Figure 2) large with four to five apical micro-setae.

Figures 1–3.

Ectinorus (Ectinorus) ineptus (Johnson, 1957) neallotype. 1: anal stylet; 2: terminal segment and unciform sclerotization; 3: spermatheca and ducti.

Spermatheca: Bulga angular (Figure 3), slightly wider than long; hilla slightly curved, twice as long as bulga; ductus obturatus and ductus spermathecae of same width, the d. s. twice as long as the d. o. Typically, for this group, ductus bursa appears as a curved “ε”.

Dimensions (slide-mounted insects in personal collection): males 2.1–2.8 mm (holotype 2.7), females 2.5–2.8 mm.

Specificity: Auliscomys Osgood, 1915 (Rodentia – Cricetidae – Sigmodontinae) must be the elective type host-genus; it is taxonomically very close to Phyllotis Waterhouse, 1837.

Ectinorus (Ectinorus) insignis Beaucournu & González-Acuña n. sp.

urn:lsid:zoobank.org:act:31EF7F63-C723-4961-8628-7548325BEAB6

Authorship: Note that the authors of the new taxon are different from the authors of this paper; Article 50.1 and Recommendation 50A of International Code of Zoological Nomenclature.

Material examined: Holotype male on Eligmodontia puerulus (Philippi, 1896) (Rodentia – Cricetidae – Sigmodontinae), Toconao (23° 07′ S–68° 00′ O), altitude 2469 m, (El Loa), Chile, X/11/2011. Allotype female in syntopy with the holotype (D. González-Acuña rec.).

Material deposited: Types in the collection of first author (Collection JCB, currently in Faculty of Medicine, Rennes), for ultimate deposition in Muséum National d’Histoire Naturelle, Paris (MNHN).

Etymology: from the Latin word insignis derived from in signum “showing a particular character”, this in reference to the male genitalia.

Description

Ectinorus insignis n. sp. has a well-developed processus basimeris ventralis and belongs to the “hapalus-group” (Smit, 1987) [36]. From a dichotomous view, this character is not chosen by this author who uses the presence or absence of spinelets on tergum I.

Head capsule: frontal tubercle not very prominently situated a short distance from the labial angle. Maxillary and labial palps extend slightly beyond the apex of trochanter. Segments I and III of maxillary palpus are the same length but slightly shorter than II or IV. Labial palpus with eight segments, including basal segment. Pre-antennal seta of male well developed, but very thin and small in female. Three pre-ocular setae, the ventral one longer as typical. Eye pigmented, indented and with sclerotized “bridge” near the gena. One post-ocular seta, three marginal setae and one very small spiniform seta on angle between gena and antennal fossa. three occipital rows with one, one and four setae in the male but only two rows in the female of one and four setae.

Thorax: prothorax with one marginal row of six well-developed setae and one anterior row of three. Male with single pseudoseta, female with two. Mesothorax with one marginal row of four long setae and anteriorly three to four small ones; one row of pseudosetae, 13 in the male, 12 in the female. Metathorax with one marginal row of four well-developed setae. Metepimeron with four setae and the spiracle shaped like a “nightcap”. Femur I: femoral guard setae very unequal in length (internal seta length and thickness twice that of the external one). Femur III: internal femoral-tibia guard seta four times as long as the external one but only slightly thicker. Tibia III with six dorsal notches bearing setae, the second, fourth and sixth with at least one long slender seta. First tarsomere III with a long apical seta as long as this segment. Second tarsomere III with a tuft of three long setae, the longest extending beyond the apex of the distitarsomere in the male and half that length in the female.

Abdomen (unmodified segments): male tergum I without spinelet while the female has one or two small spinelets. Terga II–VII, one principal row of six setae, the most ventral at level of spiracle in the male but below the spiracle in the female. Spiracles practically round in the male whereas shaped like a «nightcap» in the female. One ante-sensilial seta; marginal in the female, as typical. Sensilium typical with 17–18 pits. Sternum II with two thin marginal setae; three thin lateral setae (male) or 16–18 (female). Striarium absent. Sterna III and IV with row of three thin setae (male) or two to three (female). Sterna V–VII with one row of two thin setae (male) or respectively three, five and six long, thin and gently curved setae (female).

Abdomen (male genital segments): Sternum VIII (Figure 4) with inferior-distal lobe as in numerous species of Ectinorus. Lobe, circular with two small curved setae on its posterior margin, linked to the segment by a well-sclerotized connection (in lateral aspect). Segment IX (Figure 5) with very long processus basimeris ventralis appears longer due to lack of setae on posterior-margin of basimere.

Figures 4–10.

Ectinorus (Ectinorus) insignis Beaucournu & González-Acuña n. sp. Holotype, 4: sternum VIII; 5: segment IX; 6: processus telomeris; 7: phallosome apex – Allotype; 8: sternum VII; 9: unciform sclerotization, spermatheca and ducti.; 10: anal stylet.

Processus basimeris or Basimere: anterior and posterior edges subequal. About 10 thin setae on anterior and external borders. Margin of posterior border with one row of nine setae (slightly thicker than the drawing). Below this row, border is hyaline and naked. Inferior margin with thin setae. Ventral margin of manubrium forms a straight line visible but not as marked as in E. ineptus. Processus basimeris ventralis narrow, relatively long, with an acuminate apex; apex is anteriorly folded up as in a penknife. External part with three long and six shorter setae on its lateral margin. Internal part with only small setae. Dorsal margin of p.b.v., with two long setae; most anterior seta at base of this process

Processus telomeris (Figure 6) or Telomere, entirely masked by basimere, not extending beyond basimere as in simonsi and nomisis. Telomere longer than broad with triangular apex posterior end forming a short forward process.

Sternum IX (Figure 5): Proximal arm broad basally, narrower ventrally; distal arm narrow, then broader with a pointed apex. One short median seta and seven long curved setae on ventral margin. Tuft of six to seven small thin setae on internal surface.

Phallosome (Figure 7): classical structure for this genus, and subgenus, but apico-ventral lobe particularly broad.

Abdomen (female genital segments): Tergum VIII with four to five small setae above spiracle, spiracle elongated and vermiform; laterally about 12 small setae. Unciform sclerotization visible above spermatheca, on tergum VIII, shaped like a “horseshoe” and slightly pigmented (Figures 8 and 9).

Sternum VII (Figure 8): posterior margin convex on ventral half with five long setae remote margin; no other seta. Anal stylet (Figure 10) cylindrical with one apical seta long and gently curved, and basally one very small seta. Sternum VIII longer than wide (and very different from E. ineptus in this regard).

Spermatheca (Figure 9): bulga somewhat quadrangular; area cribriformis in the middle of proximal end; hilla broad and subrectangular. Ductus bursae with shape of a cursive ε, characteristic of this species-group, for example in ineptus, convexus, hirsutus, etc.

Dimensions (slide-mounted specimens): holotype male 1.9 mm; allotype female 2.7 mm.

Diagnosis

Among the species of Ectinorus sensu stricto, E. ineptus Johnson, 1957 [19], E. hirsutus Hastriter, 2009 [15] and E. splendidus Smit, 1968 [34] show some similarities with E. insignis n. sp. These species belong to Ectinorus having marginal spinelets on tergum I, they also belong to the hapalus-group having processus basimeris ventralis. This group shares common characters with the former genus Dysmicus and the genus Ectinorus sensu stricto (Smit, 1987) [36].

A non-genital difference is noted for the first time. Ectinorus are split into two groups by Smit [36] with respect to whether or not they have spinelets on the margin of tergum I. As we have stated, E. insignis n. sp. belongs to the hapalus-group, having processus basimeris ventralis, but unlike ineptus and hirsutus the male has no spinelet on tergum I and the female has only one or two instead of the more typical, four or six. Indeed, Dysmicus Jordan, 1942 [21] is characterized by no spinelet on tergum I, seems to mirror Ectinorus, in this respect.

Segment IX: Basimere has the same general shape as in the taxa previously mentioned. However in insignis it is less compact; the posterior margin and chaetotaxy are also different. E. splendidus, hirsutus and insignis have almost the same processus basimeris ventralis (pbv), but in E. insignis it is straight and not bent at right angles as in E. hirsutus. The pbv is short, massive and inserted at the base of basimere in E. ineptus. Among E. Ectinorus, only E. splendidus and E. insignis share-related shapes of the pbv. Sternum IX is close to that of E. ineptus by its pointed apex. The lobe of sternum VIII is not present in E. splendidus. This lobe is almost circular in E. insignis and in E. ineptus. However it bears only two setae in E. insignis while there are eight or nine in E. ineptus (six or eight in our specimens); E. hirsutus also has a circular ventrally indented lobe. For E. insignis, sternum VII only has five setae adjacent from the margin, whereas there are 10–11 setae (11 and 12 in our specimens) close to the margin in E. ineptus and E. hirsutus. The sternum of E. splendidus is very different.

Females are very similar to each other (the female of E. splendidus is currently unknown) and always have unciform sclerotization.

Host specificity: At this time, Ectinorus insignis n. sp. is only known from Eligmodontia puerulus, a sigmodontine rodent, which is also a secondary host of Ectinorus ineptus, as cited above.

Unciform sclerotization

These are “sclerotised fold (or folds) on the anterior portion of tergum VIII of the female in some fleas, usually situated under sternum VII and near its margin” (Rothschild & Traub, 1971) [31], studied also by Smit (1970) [35] and Peus (1976) [30]. They were first drawn by Jordan (1936) [20] with the description of Ctenophthalmus singularis (presently C. (Ethioctenophthalmus) singularis), then quoted by Smit (1963) [33] in a “Species-group in Ctenophthalmus” and quoted again by Hopkins & Rothschild (1966) [17] in volume IV of “the Catalogue of the Rothschild collection of Fleas (Hystrichopsyllidae: Ctenophthalminae)” (now Ctenophthalmidae) and finally thoroughly described by Peus (op. cit.) always in various Ctenophthalmus Kolenati, 1856: as in the subgenera Ethioctenophthalmus Hopkins & Rothschild, 1966 (Figure 11), Euctenophthalmus Wagner, 1940 (Figure 12) and Metactenophthalmus Peus, 1976. Among Ctenophthalmus, on one side, the “sclerotisations”, as spelled by Rothschild & Traub (1971) [31], are rarely described in Ethioctenophthalmus a highly polymorphic subgenus which is undoubtedly polyphyletic, on the other hand, they are obvious in C. (E.) leptodactylus Hubbard, 1963, more discreet in C. (E.) smithersi De Meillon, 1952. However, they are always seen in Euctenophthalmus and their relatives in Metactenophthalmus.

Figures 11–25

Unciforms sclerotizations in various species (the drawing of the outline of the spermathaeca is also included for scale and site of sclerotization) – Ctenophthalmidae, Ctenophthalminae: 11: Ctenophthalmus (Ethioctenophthalmus) eximius Jordan & Rothschild, 1913, Tanzania; 12: C. (Euctenophthalmus) savii calabricus Beaucournu, Valle & Launay, 1981, Italia – Malacopsyllidae: 13: Malacopsylla grossiventris (Weyenbergh, 1879), Argentina; 14: Phthiropsylla agenoris (Rothschild, 1904), Argentina – Rhopalopsyllidae, Parapsyllinae: 15: Delostichus coxalis (Rothschild, 1909), Chile; 16: Tetrapsyllus (Tetrapsyllus) maulinus Beaucournu & Gallardo, 1978, Chile; 17: Tetrapsyllus (Heteropsyllus) satyrus Beaucournu & Torres-Mura, 1986, Chile; 18: Listronius plesiomorphus Beaucournu & Gallardo, 1991, Chile; 19: Parapsyllus senellarti Beaucournu & Rodhain, 1990, Amsterdam Island; 20: Parapsyllus nestoris antichthones Smit, 1979, Antipodes Island – Rhopalopsyllinae: 21: Rhopalopsyllus lugubris lugubris Jordan & Rothschild, 1908, French Guyana; 22: Tiamastus helicis Beaucournu & Carmen Castro, 2003, Argentina; 23: Polygenis (Gephyropsylla) klagesi klagesi (Rothschild, 1904), French Guyana; 24: Polygenis (Polygenis) rimatus (Jordan, 1932), Brasil; 25: Polygenis (Neopolygenis) pradoi (Wagner, 1937), Colombia.

As Peus [30] showed, these sclerotizations are complex. Depending on the species or genus, the structures will be more or less visible. So the large differences noted in forms are not surprising, for example between Ctenophthalmidae and Malacopsylloidea. This structure is also present in a number of other Families: for instance, slerotization is visible in two widely distributed species, Leptopsylla segnis (Schönherr, 1811) and Nosopsyllus fasciatus (Bosc d’Antic, 1800), Ceratophyllidae (respectively Leptopsyllinae and Ceratophyllinae) and shows similarities with those of Rhopalopsyllinae (Beaucournu, unpublished).

In the Superfamily Malacopsylloidea, even though they are consistent in a given species, unciform sclerotizations are random with respect to genus or other taxa. Their shape varies from one species or genus to another. In our opinion, the interest is not only in their shape but also their presence or absence. In the genus Ectinorus, Smit (1987) [36] draws an unciform sclerotization, without comment, for some species such as E. levipes (Jordan & Rothschild, 1923), E. viscachae (Wagner, 1937) (e.g., Ectinorus) and E. onychius (Jordan & Rothschild, 1923) (s. g. Ichyonus). But, this sclerotization is also visible in Ectinorus barrerai Jordan, 1939, E chilensis Lewis, 1976, E. cocyti (Rothschild, 1904), E. ixanus (Jordan, 1942), E. martini Lewis, 1976 and E. setosicornis Jordan, 1942. However, Smit [36] seems to have overlooked this structure when he studied these taxa. Thereafter, some species with the sclerotization were described after the publication of Smit’s “Monograph”. In E. uspallatae Beaucournu & Gallardo, 2005 [6], for example, reported it as a lacuna, a name proposed by Peus [30] for some of these structures. Conversely, it is lacking in E. (Panallius) galeanus (Jordan, 1939).

Unciform sclerotization exists in other Rhopalopsyllidae. Smit [36] drew it, for example, in various Parapsyllus and in Malacopsyllidae. This family is composed of two monotypic genera, Malacopsylla Weyenberg, 1879 and Phthiropsylla Wagner, 1939. Rhopalopsyllidae are relatives of Malacopsyllidae and form with them the Superfamily Malacopsylloidea.

Smit (1987) [35] and various authors (including Johnson, 1957 [19], Linardi & Guimarães, 1993 [26], 2000 [27]) have drawn them in Rhopalopsyllidae. But the discussion must extend beyond these drawings, because the authors did not seem interested in the structure. However, Linardi & Guimarães (2000) [27] point out “mancha escura” (dark spot) or “mancha clara escamiforme” (scaly bright spot) in several Rhopalopsyllinae. Those are various unciform sclerotizations.

Combining these data and ours, we report unciform sclerotization for Malacopsylloidea in Table 1.

Table 1.

Presence or absence of unciform sclerotization in the superfamily Malacopsylloidea.

Family	Subfamily	Genus	Subgenus	Species/subspecies	Presence or absence of unciform sclerotization
Malacopsyllidae		Malacopsylla Weyenberg, 1881		grossiventris 1 (Weyenberg, 1879) (Figure 13)	Presence Visible in Smit’s drawings (1987) [35]
		Phthiropsylla Wagner, 1939		agenoris 1 (Rothschild, 1904) (Figure 14)
Rhopalopsyllidae	Parapsyllinae	Delostichus Jordan, 1942		ojedoi Beaucournu & Gallardo, 2005 smiti Jameson & Fulk, 1977 [18] talis (Jordan, 1936) xenurus (Rothschild, 1914)	Females not studied or unknown Presence or absence undeterminable
				coxalis (Rothschild, 1909) (Figure 15) degus Beaucournu, Moreno & González-Acuña, 2011 [10] incisus Beaucournu & Torres-Mura, 1987 [12] octomyos Jordan, 1942 [22] phyllotis Johnson, 1957 [19]	Presence Sometimes difficult to see in some coxalis and incisus, but these specimens may have been overcleared
		Tetrapsyllus Jordan, 1931	Phylliver Smit, 1987	bleptus (Jordan & Rothschild, 1923) Females provisionally assigned to elutus Johnson, 1957	Presence Males not studied
			Tetrapsyllus s. sto	comis Jordan, 1931 contortrix Jameson & Fulk, 1977 tristis Johnson, 1957 [19]	Females not studied Presence or absence undeterminable
				amplus (Jordan & Rothschild, 1923) corfidii (Rothschild, 1904) maulinus Beaucournu & Gallardo, 1978 (Figure 16) rhombus Smit, 1955 tantillus (Jordan & Rothschild, 1923)	Presence Sclerotization in T. rhombus is visible in Figure 117 of Smit (1987) [36]
			Heteropsyllus Beaucournu, 2002 [1]	incisus 1 Beaucournu & Torres-Mura, 1986 (Figure 17) [11]	Presence
Rhopalopsyllidae	Parapsyllinae	Ectinorus Jordan, 1942	Panallius Jordan, 1942	galeanus 1 (Jordan, 1939)	Absence
			Ectinorus s. sto.	barrerai Jordan, 1939 chilensis Lewis, 1976 [25] cocyti (Rothschild, 1904) curvatus Beaucournu & Gallardo, 1991 gallardoi 2 Hastriter, 2001 [14] hirsutus Hastriter, 2009 ineptus Johnson, 1957 (Figure 1) insignis n. sp. (Figure 10) ixanus (Jordan, 1942) levipes (Jordan & Rothschild, 1923) martini Lewis, 1976 [25] nomisis Smit, 1987 pilosus Beaucournu & Carmen Castro, 2002 setosicornis Jordan, 1942 (E. simonsi Rothschild, 1904, not examined but included with nomisis (see above)) splendidus Smit, 1968 uspallatae Beaucournu & Gallardo, 2002 viscachae (Wagner, 1931)	Presence Female of lagidium Beaucournu & Gallardo, 2005 [8] is unknown Female of mimacydis Beaucournu & Gallardo, 2004 [7] is unknown
			Ichyonus Smit, 1987	onychius onychius (Jordan & Rothschild, 1923)	Presence Visible in Smit’s drawings (1987) [35]
		Eritranis Jordan, 1942		andricus 1 (Jordan, 1939)	Presence
		Listronius Jordan, 1942		fortis (Jordan & Rothschild, 1923) plesiomorphus Beaucournu & Gallardo, 1991 (Figure 18) robertsianus 2 (Jordan, 1938) ulus (Jordan & Rothschild, 1923)	Presence
		Parapsyllus Enderlein, 1903		cardinis* Dunnet, 1961 dacunhai De Meillon, 1952 heardi De Meillon, 1952 humboldti Jordan, 1942 jacksoni* Smit, 1965 longicornis* (Enderlein, 1901) lynnae lynnae* Smit, 1965 lynnae alynnae* Smit, 1979 magellanicus magellanicus* Jordan, 1938 nestoris nestoris* Smit, 1965 nestoris antichtones Smit, 1979 (Figure 20) (cardinis group) senellarti Beaucournu & Rodhain, 1990 (Figure 19) struthophilus Smit, 1979 valedictus* Smit, 1979 (longicornis group)	Presence *indicate Smit’s drawings (1965, 1979, 1987) with unciform sclerotization For P. longicornis, note that Christchurch’s females (from Amsterdam Island) clearly show unciform sclerotization while P. longicornis from Isla Magdalena (Southern Chile) do not or barely show it.
	Rhopalopsyllinae	Rhopalopsyllus Baker, 1905		australis australis (Rothschild, 1904) australis tamoyus Jordan & Rothschild, 1923* [24] lugubris lugubris Jordan & Rothschild, 1908 (Figure 21)* lutzi lutzi (Baker, 1904)* saevus Jordan & Rothschild, 1923 [24]	Presence *indicate a “mancha escura” reported by Linardi & Guimarães (2000) [27]
				garbei (Guimarães, 1940)	No or little sclerotization
		Tiamastus, Jordan, 1939		callens (Jordan & Rothschild, 1923) [24] cavicola (Weyenbergh, 1881) deflatus Smit, 1987* gallardoi Beaucournu & Kelt, 1990 [9] helicis Beaucournu & Carmen Castro, 2003 (3) (Figure 22) palpalis (Rothschild, 1911)* plesius Jordan, 1942* subtilis (Jordan & Rothschild, 1923) [24] tortuosus Beaucournu & Carmen Castro, 2003 [3]	Presence *indicate Smit’s drawings (1987) [36] with unciform sclerotization
		Scolopsyllus Méndez, 1968		colombianus 1, 2 Méndez, 1968 [29]	Neither the original description nor Smit show sclerotisation
		Hechtiella Barrera, 19522		nitidus 2 Johnson, 1957	Presence on Figure 190 in Linardi & Guimarães (2000) [27]
				lakoi 2 Guimarães, 1948 lopesi 2 Guimarães & Linardi, 1993
		Neotropsylla Linardi & Guimarães, 1993 [26]		guimaraesi 1, 2 Linardi, 1978
		Polygenis Jordan, 1939	Gephyropsylla Barrera, 1952	klagesi klagesi (Rothschild, 1904) (Figure 23) klagesi rangeli 2 Smit, 1987 klagesi samuelis (Jordan & Rothschild, 1923) [24]
			Ayeshaepsylla Smit, 1987	thurmanni 2 Johnson, 1957	Presence in all taxa studiedSmit (op. cit.) [36] shows no sclerotization for any of these species or subspecies
			Neopolygenis Linardi & Guimarães, 1993 [26]	atopus (Jordan & Rothschild, 1922) dentei 2 Guimarães, 1947 frustratus Johnson, 1957 massoiai Del Ponte, 1967 pradoi (Wagner, 1937) (Figure 25) pygaerus 2 (Wagner, 1937)	Linardi & Guimarães (op. cit.) [27] note a “mancha escura” for: P. klagesi klagesi P. adelus P. atopus And a “mancha clara escamiforme” for P. tripusThese are unciform sclerotizations recognizable on photographs (Figure 171, 211, 302 and 330 in Linardi & Guimarães, 2000) [27]
Rhopalopsyllidae	Rhopalopsyllinae	Polygenis Jordan, 1939	Polygenis Jordan, 1939	acodontis 2 (Jordan & Rothschild, 1923) [24] adelus 2 (Jordan & Rothschild, 1923) [24] adocetus 2 Traub, 1950 axius axius (Jordan & Rothschild, 1923) [24] axius pessoai 2 Guimarães, 1956 axius proximus 2 Guimarães, 1948 bolhsi bolhsi (Wagner, 1901) bolhsi jordani (Da Costa Lima, 1937) brachinus Jordan, 1950 caucensis 2 Méndez, 1977 delpontei Méndez, 1977 dendrobius 2 (Wagner, 1939) dunni (Jordan & Rothschild, 1922) floridanus 2 Johnson & Layne, 1961 gwyni (Fox, 1914) guimaraesi 2 Linardi, 1978 hopkinsi 2 Méndez, 1977 impavidus 2 Johnson, 1957 litargus 2 (Jordan & Rothschild, 1923) [24] litus 2 (Jordan & Rothschild, 1908) martinezbaezi 2 Vargas, 1951 occidentalis occidentalis 2 (Cuhna, 1914) [13] occidentalis steganus 2 (Jordan & Rothschild, 1923) [24] odiosus 2 Smit, 1958 platensis (Jordan & Rothschild, 1908) peronis 2 (Jordan & Rothschild, 1923) puelche 2 (Del Ponte, 1963) rimatus (Jordan, 1932) (Figure 24) roberti beebei (Fox, 1947) roberti roberti 2 (Rothschild, 1905) roberti tripopsis 2 Guimarães, 1948 rozeboomi 2 Vargas, 1952 trapidoi mendezi 2 Smit, 1987 trapidoi trapidoi 2 Méndez, 1977 tripus 2 (Jordan, 1933) vazquezi 2 Vargas, 1951	Presence in all taxa studied Smit (op. cit.)[35] shows no sclerotization for any of these species or subspeciesLinardi & Guimarães [27] note a “mancha escura” for: P. klagesi klagesi P. adelus P. atopus And a “mancha clara escamiforme” for P. tripus These are unciform sclerotizations recognizable on photographs (Figure 171, 211, 302 and 330 in Linardi & Guimarães, 2000) [27]

Monotypic genus.

Taxa not studied.

colombianus Méndez, 1968 [29]

guimaraesi Linardi, 1978

In conclusion, there is unciform sclerotization for the two genera of Malacopsyllidae. In Rhopalopsyllidae – Parapsyllinae, it is present for every species of Delostichus, Tetrapsyllus, Ectinorus, Eritranis and Listronius studied. However, curiously, the enigmatic Ectinorus (Panallius) galeanus does not have it. Finally, it seems to be present in all species and subspecies of the subfamily Rhopalopsyllinae in the Rhopalopsyllidae. This structure varies in size and shape depending on the species but its presence can help species classification. We did not aim to search for the slerotization in all Siphonaptera, but hopefully by this note, we will draw attention to this structure by other researchers. Indeed the structure appears to have taxonomic value in the classification of particular subgenera within the genus Ctenophthalmus. Females, for example in Rhopalopsyllidae, are difficult to identify. Therefore, the presence or absence of sclerotization must be well figured or at least reported to aid in identification.