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The chewing lice (Insecta: Phthiraptera: Amblycera and Ischnocera) of Mexico have been little studied and many publications include isolated records. This paper summarizes current knowledge of chewing lice recorded from Mexico resulting from an exhaustive search of the literature published from 1866 to 2017. We found 342 louse species associated with 206 bird and 28 mammal species. As a result, we provide a checklist of the chewing lice recorded from Mexico, including a host-parasite list and their geographical distribution within the country.

Family Fregatidae Fregata magnificens Mathews Colpocephalum spineum Pectinopygus fregatiphagus, Published as part of S��nchez-Montes, Sokani, Colunga-Salas, Pablo, ��lvarez-Castillo, Luc��a, Guzm��n-Cornejo, Carmen & Montiel-Parra, Griselda, 2018, Chewing lice (Insecta: Phthiraptera) associated with vertebrates in Mexico, pp. 1-109 in Zootaxa 4372 (1) on page 82, DOI: 10.11646/zootaxa.4372.1.1, http://zenodo.org/record/3098143

Family Fregatidae Fregata magnificens Mathews Colpocephalum spineum Pectinopygus fregatiphagus

Limnofregata hasegawai, new species (Figs. 5–7) Holotype: GMNH PV 170, skull and mandible with quadrates, lacrimals, palatines, atlas, axis, and cervical vertebrae 3–7 (Fig. 5). Color brownish black. The skull is preserved in right dorso­lateral view with the mandible and right quadrate still in articulation. The left quadrate has been displaced and lies upside­down, with the mandibular articulation visible along the edge of the left orbit. The lacrimals are still nearly in articulation with the frontals, the right palatine is exposed but neither pterygoid is visible. Locality: Smith Hollow Quarry, Lewis Ranch (Locality B of Grande and Buchheim, 1994), Lincoln County, Wyoming. Horizon: Said to have been collected " 2–3 m " from Referred Specimen 1 of L. azygosterno n (GMNH PV 167). Presumably this distance is intended as vertical as the matrix and the coloration of the bone (blackish vs. brown) are different. Fossil Butte Member of the Green River Formation, late early Eocene. Etymology: To our esteemed colleague and friend Yoshikazu Hasegawa of the Gunma Museum of Natural History in recognition of his many contributions to vertebrate paleontology. Diagnosis: Much larger than L. azygosternon. Rostrum proportionately much longer, 1.6 times as long as the cranium, vs. 1.4 in L. azygosternon. Measurements (mm) of holotype: Measurements that are comparable to those published for Limnofregata azygosternon appear in Table 2. Additional measurements are: Cranium: width at nasofrontal hinge, 16.8. Lacrimal: length, 17.1. Quadrate: length of articulation with mandible (left), 15.8. Atlas: width 11.9. Axis: width 13.1. Cervical vertebra 3: length, 16.1. Cervical vertebra 5: length, 22.8. Paratype 1: FMNH PA 719, complete skull and mandible with left quadrate, both lacrimals and pterygoids, assorted sclerotic plates, atlas, axis, and cervical vertebra 3 (Fig. 6). The skull is preserved in dorsal and partial left lateral view with the entire left quadratojugal bar visible. The ventral surface of the mandible can be seen in its entirety. Locality: Thomson Ranch (Locality H of Grande and Buchheim, 1994), ca. 14 km W of Kemmerer, Lincoln County, Wyoming. Horizon: F­ 2 facies, Fossil Butte Member of the Green River Formation, late early Eocene. Measurements (mm): The following are in addition to those in Table 2. Cranium: width at nasofrontal hinge, ca. 22. Pterygoid: length, 23.1. Quadrate: depth, 20.6; width of mandibular articulation, 15.7; length of otic process along dorsal edge, 12.2. Lacrimal: length, 18.1; depth including descending process, 19.0; depth of descending process, 12.5; greatest depth of corpus at posterior end 7.9. Mandible: posterior width (distance between external margins of articulations), 50.7; width of articulation, 17.3. Atlas: width and depth, 12.6 X 10.9. Paratype 2: BMS E 25336, pelvis with associated right and left femora and tibiotarsi, the first 5 free caudal vertebrae, and 10 presacral vertebrae (Fig. 7). Collected by Verl and Rick Hebdon and acquired by the Buffalo Museum of Science in 1982. Locality: Warfield Fossil Quarries at Warfield Springs (NW 1 / 4, Sec. 5, T 19 N, R 117 W; locality K of Grande and Buchheim, 1994), ca. 14 km SW of Kemmerer, Lincoln County, Wyoming. Horizon: F­ 2 facies equivalent, Fossil Butte Member of the Green River Formation, late early Eocene. The specimen is from a 1.5 m thick layer of precipitated limestone overlying a 30–45 cm thick layer of bluish shale. Measurements: See Table 1. Description: As in L. azygosternon the nostril is long and open, the posteriormost corner being about 1 cm anterior to the naso­frontal hinge. The anterior margin can now be clearly discerned as extending nearly to the premaxillary symphysis. The configuration of the quadrate in ventral view in Limnofregata shows some distinctive features. In Phaethon the medial and lateral condyles are in the same transverse plane with practically no development of a posterior condyle. In Fregata, the medial condyle extends posterolaterally as a more constricted crest, which then bends at nearly a right angle to the lateral condyle, the bend forming a rather weak posterior condyle with a deep depression anterior to it. The configuration in the Sulidae is more complex, with the posterior condyle completely separate from the medial condyle. Limnofregata is most similar to Fregata, with the medial condyle forming a long rolling crest with only an incipient posterior condyle and the lateral condyle is only slightly angled off of the long axis of the articular surface. The depression seen in Fregata is shallower in Limnofregata. The pterygoids are relatively much longer in Limnofregata than in Fregata, with slightly curved rather than straight shafts that appear to be terete, without the flattened surfaces seen in Fregata or the Sulidae. They are quite unlike the very long, straight, and slender pterygoids in Phaethon. The mandibular articulations are much heavier in Limnofregata than in Fregata, the rami much thicker and deeper, and the mandibular symphysis shorter. In the ventral view afforded by the paratypical skull of L. hasegawai, the internal processes of the articulations are shorter, thicker, and much less distinctly set off from the heavy proximal portion of the rami than in Fregata. The one visible lacrimal in the holotype of L. azygosternon is partially obscured and was misinterpreted in the original description. The entire structure is beautifully revealed in paratype FMNH PA 719 of L. hasegawai in which both lacrimals are preserved with the entire external surface exposed (Fig 6). The corpus of the bone that articulates with the cranium is much larger than in Fregata, with the posterior portion broad and truncated. There are two small pneumatic foramina in the corpus, just anterior to the descending process. The descending process is very long and slender, with a very small expansion on the posterior margin about 2 / 3 the distance to the ventral extremity. The lacrimal in Fregata is very different, with a small, nonpneumatic corpus and a greatly expanded, inflated descending process that bears a single pneumatic foramen. The postcranial specimen of L. hasegawai is not in particularly good condition, but has the pelvis preserved in dorsal view and the thoracic vertebrae are better preserved than in the holotype of Limnofregata azygosternon. These do not differ greatly from those in Fregata. The specimen confirms that the distinctive notch in the posterior margin of the innominate bone in the area of fusion of the ischim and ilium (Olson, 1977: 15) is the normal condition. As in the holotype of L. azygosternon, the pygostyle is missing, suggesting that it may have floated away with the presumably enlarged rectrices. Size in Limnofregata. The seven available specimens now assigned to L. azygosternon are remarkably homogeneous in size and those for which comparable length measurements are available are practically identical (Table 1), especially given the variation induced by crushing and other vicissitudes of preservation. In the two skulls of L. hasegawai such factors of preservation have doubtless contributed to inaccuracies in the measurements of the cranium and rostrum. For example, in the paratype FMNH PA 719, the skull is broken across the nasofrontal hinge and the anterior margin of the cranium has been shoved under the posterior margin of the rostrum perhaps as much as 6 mm. On the other hand, the length measurements of the two known mandibles of L. hasegawai and in the holotype of L. azygosternon are unequivocal. The two mandibles of L hasegawai are identical in length and are 20 % larger than in the holotype of L. azygosternon. The postcranial paratype of L hasegawai basically agrees in size, as the femur is 17 % larger and the pelvis 20 % larger than in L. azygosternon. The tibiotarsus seems disproportionately small, however, as it is only 7 % larger. The same appears to apply within modern frigatebirds, however, as the tibiotarsus of the largest species, F. m i n o r, is proportionately smaller than in F. a r i e l, the smallest species (Table 3). The differences in size between L. azygosternon and L. hasegawi are as great as between the smallest and largest individuals of modern Fregata, which comprises 5 sexually dimorphic species (females larger). Thus, on size alone, Limnofregata would have to be divided into two species­level taxa, as the differences could not be due to differences between sexes, especially as the species of Limnofregata are unlikely to have been sexually dimorphic (see discussion below). The differences in bill proportions, with L. azygosternon having proportionately a much shorter bill, would also not be expected within a single species.

Genus Limnofregata Olson, 1977 Type��species: Limnofregata azygosternon Olson, 1977. Included species: L. azygosternon Olson, 1977: L. hasegawai, new species., Published as part of Olson, Storrs L. & Matsuoka, Hiroshige, 2005, New specimens of the early Eocene frigatebird Limnofregata (Pelecaniformes: Fregatidae), with the description of a new species, pp. 1-15 in Zootaxa 1046 on page 2, DOI: 10.5281/zenodo.169909, {"references":["Olson, S. L. (1977) A Lower Eocene frigatebird from the Green River Formation of Wyoming (Pelecaniformes, Fregatidae) Smithsonian Contributions to Paleobiology, 35, 1 - 33."]}

Limnofregata azygosternon Olson, 1977 Figs. 1���4 This species was described from a nearly complete skeleton with feather impressions (holotype USNM 22753) and two paratypes consisting of most of a right wing and shoul�� der girdle (UWY 6919) and the proximal end of an ulna (USNM 243766). Here we identify four additional specimens that can also be referred to the species Limnofregata azygosternon. Referred specimen 1: GMNH PV 167, postcranial skeleton with impressions of remiges and contour feathers (Fig. 1). Bone color brown, feathers black. Collected by Tom Lindgren. The specimen consists of all elements of the appendicular skeleton except the proximal portion of the right humerus, plus the complete pectoral girdle and pelvis. It is lacking the skull and mandible, a number of ribs, and the entire pre�� and postsacral vertebral column except for three thoracic and a caudal vertebra. Very distinct carbonized impressions of feathers can be identified as primaries, secondaries, and wing coverts. Locality: Smith Hollow Quarry, Lewis Ranch (Locality B of Grande and Buchheim, 1994), Lincoln County, Wyoming. Horizon: Beds about 4 m above the ��� 18 inch layer,��� Fossil Butte Member of the Green River Formation, late early Eocene. Measurements (mm): See Table 1. Referred specimen 2: FMNH PA 723, posterior portion of an associated skeleton consisting of the last 3 thoracic vertebrae, pelvis, caudal vertebrae and pygostyle, both hindlimbs complete with all toe bones, and assorted ribs (Fig. 2). Locality: Thomson Ranch (Locality H of Grande and Buchheim, 1994), ca. 14 km W of Kemmerer, Lincoln County, Wyoming. Horizon: F�� 2 facies, Fossil Butte Member of the Green River Formation, late early Eocene. Measurements (mm): See Table 1. Referred specimen 3: FMNH PA 720, complete left wing distal to the humerus (Fig. 3). Locality: Warfield Springs (NW 1 / 4, Sec. 5, T 19 N, R 117 W; locality K of Grande and Buchheim, 1994), ca. 14 km SW of Kemmerer, Lincoln County, Wyoming, on the SE shore of Eocene Fossil Lake. Horizon: F�� 2 facies equivalent, Fossil Butte Member of the Green River Formation, late early Eocene. Measurements (mm): See Table 1. Referred specimen 4: USNM 447002, left side of cranium with quadrate and associated posterior portion of mandible (Fig 4). The base of the bill and the occiput were preserved as a mold that has now been replaced with embedding compound. Locality: Ca. 12 km NE of Fontanelle Reservoir Dam on N shore at autumn water level, Sweetwater County, Wyoming. This locality is in beds of the Eocene Lake Gosiute, whereas all the other specimens of Limnofregata to date have come from deposits in Fossil Lake. Horizon: Laney Shale Member of the Green River Formation, lower middle Eocene. Measurements (mm): Quadrate: distance from posterior margin of mandibular articulation to tip of otic process, ca. 13.4. Description: Although USNM 447002 is very fragmentary, it is in places less crushed than other skulls of the genus. It shows a well��developed ectethmoid plate extending laterally from the mesethmoid, whereas the ectethmoid is much more reduced in Fregata. It emphasizes the large size of the orbit in Limnofregata, suggesting that the birds may have been partially nocturnal or crepuscular. In GMNH PV 167, the toes of the right foot appear to be in their naturally maximal spread position, and shows the much better development of the foot in Limnofregata compared with Fregata. Assuming that the foot was totipalmate, as in all modern Pelecaniformes, the area of the webbing was on the order of 20 cm 2. In the largest and best preserved of the primary feathers in GMNH PV 167, the tip is much less attenuated and sharply pointed than in Fregata. For FMNH PA 723, new information not preserved with the holotype concerns the tail. The number of free caudal vertebrae anterior to the pygostyle appears to be seven, as in Fregata. The pygostyle of Limnofregata was previously unknown, this element being entirely missing in the holotype of L. azygosternon. It is a broad plate as in Phaethon and Fregata, and completely unlike the pygostyle in the Sulidae, which is very thick and somewhat cylindrical proximally, tapering to a long, thin spine. In Limnofregata, the ventral margin of the pygostyle (Fig. 2) is rectilinear, the dorsal margin is curved, and the tip is broadly rounded., Published as part of Olson, Storrs L. & Matsuoka, Hiroshige, 2005, New specimens of the early Eocene frigatebird Limnofregata (Pelecaniformes: Fregatidae), with the description of a new species, pp. 1-15 in Zootaxa 1046 on pages 2-6, DOI: 10.5281/zenodo.169909, {"references":["Olson, S. L. (1977) A Lower Eocene frigatebird from the Green River Formation of Wyoming (Pelecaniformes, Fregatidae) Smithsonian Contributions to Paleobiology, 35, 1 - 33.","Grande, L. & Buchheim, H. P. (1994) Paleontological and sedimentological variation in early Eocene Fossil Lake. Contributions to Geology University of Wyoming, 30, 33 - 56."]}

Four additional specimens from the Green River Formation of Wyoming are referred to the Eocene frigatebird Limnofregata azygosternon Olson, originally described from a nearly complete skeleton and two partial paratypes. Two skulls with mandibles and a partial postcranial skeleton are described as a new species, Limnofregata hasegawai, characterized by much larger size and a proportionately longer bill. One of the referred specimens of L. azygosternon is from Eocene Lake Gosiute, whereas all of the other specimens of Limnofregata are from Fossil Lake. The species of Limnofregata would have taken advantage of frequent periodic dieoffs of fish in the Green River lakes. Geological and climatic factors that may have influenced the paleoecology, distribution, and size variation in frigatebirds in the Cenozoic are reviewed.

O hábito da fregata (Fregata magnificiens) de roubar peixes do atobã marrom (Sula lecogaster) é bem conhecido na literatura, comportamento este que tem nos levado a classificar a primeira espécie como sendo cleptoparasita da segunda. Este trabalho, no entanto, mostra que a relação entre estas duas espécies é mais complexa e que o comportamento cleptoparasitãrio pode ser interpretado como sendo apenas parte de todo um processo de exploração mutua. Quando estas aves voam no encalço de embarcações pesqueiras, as fregatas localizam a fonte de alimento, sendo então seguidas pelos atobas que aparecem logo apos, apanhando os peixes mortos com muito mais eficiencia, de tal maneira que as fregatas não conseguem mais pescar, tendo que abandonar a atividade, Quando os atobas terminam sua atividade pesqueira, ou mesmo apos um numero deste ciclo, sugere-se que a fregata venha a procurar os atobas, roubando-lhes parte do pescado, fenomeno este bem descrito na literatura. Esta observação conduz a uma revisão da interpretação acerca da relação entre estas espécies, até então, considerada como cleptoparasitismo para a de exploração mútua. The frigatebird (Fregata magnificiens) often is referred to as a kleptoparasite of the brown booby (Sula lecogaster). This paper however, shows that the relationship between these species is more complex and that kleptoparasitism can be interpreted as just a part of a whole process of mutual exploration. When these birds fly behind fishing boats, F. magnificens locates the food source first and S. lecogaster then moves in and catches fish more efficiently in such a way that the frigatebird can no longer fish and has to fly away. When the fishing period is over then the frigatebird is thought to approach the brown boobies in order to get fish from them, as described in the literature, This leads to a revision on the interpretation about the relation between these species, from that of kleptoparasitism to this of mutual exploration.

The frigatebird (Fregata magnificiens) often is referred to as a kleptoparasite of the brown booby (Sula lecogaster). This paper however, shows that the relationship between these species is more complex and that kleptoparasitism can be interpreted as just a part of a whole process of mutual exploration. When these birds fly behind fishing boats, F. magnificens locates the food source first and S. lecogaster then moves in and catches fish more efficiently in such a way that the frigatebird can no longer fish and has to fly away. When the fishing period is over then the frigatebird is thought to approach the brown boobies in order to get fish from them, as described in the literature, This leads to a revision on the interpretation about the relation between these species, from that of kleptoparasitism to this of mutual exploration. O habito da fregata (Fregata magnificiens) de roubar peixes do atobã marrom (Sula lecogaster) é bem conhecido na literatura, comportamento este que tem nos levado a classificar a primeira espécie como sendo cleptoparasita da segunda. Este trabalho, no entanto, mostra que a relação entre estas duas espécies é mais complexa e que o comportamento cleptoparasitãrio pode ser interpretado como sendo apenas parte de todo um processo de exploração mutua. Quando estas aves voam no encalço de embarcações pesqueiras, as fregatas localizam a fonte de alimento, sendo então seguidas pelos atobas que aparecem logo apos, apanhando os peixes mortos com muito mais eficiencia, de tal maneira que as fregatas não conseguem mais pescar, tendo que abandonar a atividade, Quando os atobas terminam sua atividade pesqueira, ou mesmo apos um numero deste ciclo, sugere-se que a fregata venha a procurar os atobas, roubando-lhes parte do pescado, fenomeno este bem descrito na literatura. Esta observação conduz a uma revisão da interpretação acerca da relação entre estas espécies, até então, considerada como cleptoparasitismo para a de exploração mutua.