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Data from: Multispecies genetic structure and hybridization in the Betula genus across Eurasia
Boreal and cool temperate forests are the major land cover of northern Eurasia, and information about continental-scale genetic structure and past demographic history of forest species is important from an evolutionary perspective and has conservation implications. However, although many population genetic studies of forest tree species have been conducted in Europe or Eastern Asia, continental-scale genetic structure and past demographic history remain poorly known. Here, we focus on the birch genus Betula, which is commonly distributed in boreal and cool temperate forests, and examine 129 populations of two tetraploid and four diploid species collected from Iceland to Japan. All individuals were genotyped at seven to 18 nuclear simple sequence repeats (nSSRs). Pairwise math formula among the six species ranged from 0.285 to 0.903, and genetic differentiation among them was clear. structure analysis suggested that Betula pubescens is an allotetraploid and one of the parental species was Betula pendula. In both species pairs of B. pendula and B. plathyphylla, and B. pubescens and B. ermanii, genetic diversity was highest in central Siberia. A hybrid zone was detected around Lake Baikal for eastern and western species pairs regardless of ploidy level. Approximate Bayesian computation suggested that the divergence of B. pendula and B. platyphylla occurred around the beginning of the last ice age (36 300 years BP, 95% CI: 15 330–92 700) and hybridization between them was inferred to have occurred after the last glacial maximum (1614 years BP, 95% CI: 561–4710), with B. pendula providing a higher contribution to hybrids.
FIGURE 10 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 10. Neohemigaster antropovi sp. nov., female holotype; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
FIGURE 9 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 9. Neohemigaster ussurica (Korneyev, 2001), male; a—habitus, lateral view, b—head and thorax, dorsal view; chead, lateral view; d—head, anterior view; e—wing.
FIGURE 8 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 8. Neohemigaster tetralineata sp. nov., male holotype; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
FIGURE 7 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 7. Neohemigaster angustifrons sp. nov., male holotype; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
FIGURE 6. Neohemigaster eurysterna Hendel, 1914 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 6. Neohemigaster eurysterna Hendel, 1914, male; a—habitus, lateral view, b—head and thorax, dorsal view; chead, lateral view; d—head, anterior view; e—wing.
FIGURE 5. Pterogenia luctuosa Hendel, 1914 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 5. Pterogenia luctuosa Hendel, 1914, male; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
FIGURE 4. a in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 4. a—Pterogenia hamifera Frey, 1930, wing (from Frey's figure 3, Taf. 1); b—Neohemigaster minuspicta Hennig, 1940, wing (Typus); c—Neohemigaster flavopicta Hennig, 1940, wing (Typus); d— Neohemigaster ornata Hennig, 1940, wing (Typus).
FIGURE 3 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 3. Pterogenia tenebrica sp. nov., female holotype; a—habitus, lateral view, b—head and thorax, dorsal view; chead, lateral view; d—head, anterior view; e—wing.
FIGURE 2. Pterogenia hologaster Hendel, 1914 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 2. Pterogenia hologaster Hendel, 1914, male; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
FIGURE 1. Neohemigaster rectivena Enderlein, 1924 in Neohemigaster Malloch, 1939 and Pterogenia Bigot, 1859 (Diptera: Platystomatidae) from eastern Eurasia, with the description of four new species
FIGURE 1. Neohemigaster rectivena Enderlein, 1924, male type; a—habitus, lateral view, b—head and thorax, dorsal view; c—head, lateral view; d—head, anterior view; e—wing.
Fig. 5 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 5. Forelimb long bones. Shown are the humerus, radius, and ulna of the holotype (IPS58443) of Pliobates cataloniae gen. et sp. nov. (A to E) Partial left humerus in medial (A), posterior (B), lateral (C), anterior (D), and distal (E) views. (F to K) Left radius in medial (F), posterior (G), lateral (H), anterior (I), proximal (J), and distal (K) views. (L to O) Proximal half of the left ulna in medial (L), posterior (M), lateral (N), and anterior (O) views. (P to T) Distal fragment of the left ulna in medial (P), posterior (Q), lateral (R), anterior (S), and distal (T) views.
Fig. 1 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 1. Cranium and dentition. (A to C) Cranium of the holotype (IPS58443) of Pliobates cataloniae gen. et sp. nov.The main cranial fragments, including the basicranium and the right palate, are shown in basal view (A); details of the right palatal fragment are shown in left-lateral (B) and right-lateral (C) views. (D) Detail of the right postcanine teeth, in occlusal view (mesial is to the right).
Fig. 4 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 4. Basicranial morphology. (A) Drawing of the left basicranium of the holotype (IPS58443) of Pliobates cataloniae gen. et sp. nov., as preserved in ventral view.The jugular foramen appears artifactually larger because of the displacement of the temporal and occipital portions along the occipitotemporal suture (corrected in the reconstruction in Fig. 3). The course of the carotid canal is shown with a dashed line, based on CT images. AE, articular eminence; CAF, carotid foramen; COF, condylar fossa; EA, Eustachian aperture; EAM, external auditory meatus; EP, Eustachian process; ET, ectotympanic; FM, foramen magnum; FO, foramen ovale; GF, glenoid fossa; JF, jugular foramen; OC, occipital condyle; OTS, occipitotemporal suture; PGP, postglenoid process. (B to D) Drawings of comparable views (not to scale) in Hylobates sp. (B), Proconsul heseloni KNM RU 2036 [(C), reversed], and Victoriapithecus macinnesi KNM MB 29100a (D) (KNM, Kenyon National Museums; RU, Rusinga; MB, Maboko). Arrows denote the V-shaped, incompletely ossified ventral terminal tip of the tubular ectotympanic in the extinct taxa. [Artwork by M. Palmero]
Fig. 7 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 7. Carpal bones. Line drawings of carpal bones in Pliobates cata- loniae gen. et sp. nov. (IPS58443) are shown with those of selected anthropoid genera for comparison. (A to E) Left capitate, in radial (top) and proximal (bottom) views, of Cer- copithecus aethiops (A), Ateles paniscus (B), Pierolapithecus cat- alaunicus (C), Hylobates lar (D), and P. cataloniae (E); gray shading denotes articular areas for the second metacar- pals, and cross-hatching denotes those for the third metacarpal. (F to J) Left hamate, in radial (top) and ulnar (bottom) views, of C. aethiops (F), A. paniscus (G), Pi. catalaunicus (H), H. lar (I), and P. catalo- niae (J). (K to O) Left triquetrum, in proximo- medial (top) and distal (bottom) views, of C. aethiops (K), A. paniscus (L), Pi. catalaunicus (M), H. lar (N), and P. cataloniae (O). Drawings are not to scale.
Fig. 3 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 3. Cranial reconstruction. Virtual reconstruction of the holotype (IPS58443) cranium of Pliobates cataloniae gen. et sp. nov., including mirrored fragments, in frontal (A), lateral (B), posterior (C), basal (D), and superior (E) views. Further details are given in fig. S3 and the methods in the text.
Fig. 6 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 6. Elbow and wrist morphology. The most diagnostic features of the elbow and wrist joints of Pliobates cataloniae gen. et sp. nov. (IPS58443), denoted by arrows in drawings of the distal humerus, proximal radius, and distal ulna, are shown with those of selected extant and extinct anthropoids for comparison. (A to D) Anterior (top) and distal (bottom) views of the distal humerus in P. cata- loniae (A), Epipliopithecus vindobonensis Individual I [(B), reversed], Dendro- pithecus? sp. KNM MO 17022A (C) (MO, Moruorot), and Hylobates moloch (D). (E to H) Views perpendicular to the radial tuberosity (top) and proximal view (bottom) of the proximal radius in P. cataloniae (E), E. vindobonensis Individual I (F), Simiolus enjiessi KNM MO 63 [(G), reversed)], and H. moloch (H). (I to M) Medial (top) and distal (bottom) views of the distal ulna in P. cataloniae (I), E. vindobonensis Indi- vidual I (J), H. moloch (K), Ateles paniscus (L), and Cercopithecus aethiops (M). 1, absence of entepicondylar foramen; 2, absence of capitular tail; 3, lack of spool-shaped trochlea; 4, well-developed beveled surface for the zona conoidea; 5, small and flat area in the radial head; 6, ulnar fovea; 7, two-faceted, expanded semilunar articular surface in the ulnar head. Specimens are shown as if from the left side and are not to scale. [Artwork by M. Palmero]
Fig. 2 in Miocene small-bodied ape from Eurasia sheds light on hominoid evolution
Fig. 2. Results of the dental microwear analyses. (A) Pitting incidence (%) of Pliobates, the extant comparative sample, and pliopithecoids and extinct hominoids from Europe and Turkey. (B) Bivariate plot of striation breadth versus pitting incidence. (C) Bivariate plot of the first two canonical axes delivered by the canonical variates analysis, based on three distinct, broad dietary groups: folivores, mixed feeders and frugivores, and hard-object feeders. Colored polygons in (B) and ellipses in (C) illustrate the variability of extant dietary categories. Small black symbols denote the comparative sample of extant anthropoids, whereas large black symbols represent the centroids of each dietary category. Different symbols are employed to distinguish the various extinct species; results for Iberian hominoids and pliopithecoids are shown in red and blue, respectively, whereas those from other localities are shown in yellow and green, respectively.
Subspecies and Distribution. M. n. nivalis Linnaeus, 1766 — China, North and South Korea, Mongolia, Russia, Taiwan, and Scandinavia. M. n. allegheniensis Rhoads, 1900 — NE USA (Allegheny Mts W to Wisconsin). M. n. boccamela Bechstein, 1800 — Corsica, Italy, Portugal, Sardinia, Sicily, and Spain. M. n. campestris Jackson, 1913 — C Great Plains, USA. M. n. eskimo Stone, 1900 — Alaska and NW Canada (Yukon). M. n. namiyer Kuroda, 1921 — Japan and the Kurile Is. M. n. numidica Pucheran, 1855 — N Africa. M. n. rnixosa Bangs, 1896 — Canada and N Great Plains of USA. M. n. vulgaris Erxleben, 1777 = W & C Europe and most of C Eurasia. Introduced to New Zealand, Malta, Crete, the Azores Is, and apparently also Sao Tome I. in Mustelidae
Subspecies and Distribution. M. n. nivalis Linnaeus, 1766 — China, North and South Korea, Mongolia, Russia, Taiwan, and Scandinavia. M. n. allegheniensis Rhoads, 1900 — NE USA (Allegheny Mts W to Wisconsin). M. n. boccamela Bechstein, 1800 — Corsica, Italy, Portugal, Sardinia, Sicily, and Spain. M. n. campestris Jackson, 1913 — C Great Plains, USA. M. n. eskimo Stone, 1900 — Alaska and NW Canada (Yukon). M. n. namiyer Kuroda, 1921 — Japan and the Kurile Is. M. n. numidica Pucheran, 1855 — N Africa. M. n. rnixosa Bangs, 1896 — Canada and N Great Plains of USA. M. n. vulgaris Erxleben, 1777 = W & C Europe and most of C Eurasia. Introduced to New Zealand, Malta, Crete, the Azores Is, and apparently also Sao Tome I.
Subspecies and Distribution. S. s. scrofa Linnaeus, 1758 — W Europe, from Denmark, Germany, Poland, and Czech Republic to N Italy and N Iberian Peninsula; possibly also Albania. The taxonomic status of animals in Austria, Switzerland, Slovenia, and Slovakia is unclear but presumably these populations are included in scrofa, as are the populations of Sweden, Finland, and the Baltic states. However, restocking of once depleted populations, for example in Italy, has likely involved the introduction and mixing of this subspecies with other subspecies, such as attila. S. s. affinis Gray, 1847 — S India and Sri Lanka. S. s. algirus Loche, 1867 — Tunisia, Algeria, and Morocco, on the coastal side of the mountains or in the low montane areas. S. s. attila Thomas, 1912 — Hungary, Ukraine, C & S Belarus, Romania, Moldova, and S Russia towards the N flank of the Caucasus, but not including the Transcaucasian countries of Georgia, Armenia, and Azerbaijan. The range possibly extends as far S as the Mesopotamian Delta in Iraq, in which case it would likely include W & SW Iran, and possibly E Turkey and Syria, where it borders with lybicus. Such a range could not be easily reconciled with a statement by Groves that "the difference between pigs from N and S of the Caucasus is quite striking; Transcaucasian boars are certainly not attila." This subspecies may also extend into C Asia and include Kazakhstan, Uzbekistan, and Turkmenistan, but no data exist to support this. S. s. baeticus Thomas, 1912 — originally described from Coto Donana, S Spain, and later merged with meridionalis; also S Portugal. Unless evidence is found that these Italian and Iberian populations are the relics of a much larger formerly contiguous range, this subspecies should be kept as distinct. S. s. coreanus Heude, 1897 — Korean Peninsula. S. s. eristatus Wagner, 1839 — Himalayas S to C India and E to Indochina (N of the Kra Isthmus). S. s. davidi Groves, 1981 — the arid zone from E Iran to Gujarat, including Pakistan and NW India, and perhaps N to Tajikistan. S. s. leucomystax Temminck, 1842 — main Is ofJapan (Honshu, Shikoku, Kyushu, Nakadori, Hiburijima, Tojima, Kushima, and other smaller Is). S. s. lybicus Gray, 1868 — Bulgaria, Greece, Turkey, Syria, Jordan, Israel, Palestine, in the past also in Lybia, and Egypt. The former Yugoslavia was included in its range, which would suggest that now Slovenia, Serbia, Croatia, Bosnia and Herzegovina, Montenegro, and Kosovo are within the range of this subspecies, although the exact boundaries are unclear. Pigs from Albania have been assigned to S. s. scrofa. S. s. majori De Beaux & Festa, 1927 — C & S Italian Peninsula. S. s. menidionalis Forsyth Major, 1882 — Corsica and Sardinia, with the proviso that the two populations are very likely to be introduced or feral. S. s. moupinensis Milne-Edwards, 1871 — China, S to Vietnam and W to Sichuan. S. s. nigripes Blanford, 1875 — the flanks of the Tianshan mountains in Kyrgyzstan and NW China (Xinjiang). An animal photographed in NE Iran (Golestan) looked like this subspecies. S. s. nukiuanus Kuroda, 1924 — Iriomote, Ishigaki, Okinawa, Tokunoshima, Amamioshima, and Kakerome Is in the Ryukyu chain in extreme S Japan, though some of these populations have hybridized with introduced domesticates. S. s. sibiricus Staffe, 1922 — Mongolia and Transbaikal (S & E of Lake Baikal). S. s. tawvanus Swinhoe, 1863 — Taiwan. S. s. ussuricus Heude, 1888 — far E Russia and the Manchurian region (China). Korean populations were previously included in this subspecies, but based on new evidence, the Korean taxon seems more similar to moupinensis. S. s. vittatus Boie, 1828 — Malay Peninsula, S of the Isthmus of Kra, the offshore islands of Terutai and Langkawi, Sumatra, Riau Archipelago, Java, Bali, and a range of smaller islands around these, including Babi, Bakong, Batam, Bawean, Bengkalis, Bintan, Bulan, Bunguran, Cuyo, Deli, Durian, Enggano, Galang, Jambongan, Karimon (Riau Is), Kundur, Lagong, Laut, Lingga, Lingung, Mapor, Moro Kecil, North Pagai, Nias, Panaitan, Payong, Penang, Pinie, Rupat, Siantan, Siberut, Simeulue, Singkep, Sugi, Sugi Bawa, Telibon, Tinggi, Tuangku, and the Tambelan Is. This species was originally present from the British Is in the extreme W, through Eurasia from S Scandinavia to S Siberia, extending as far E as Korea and Japan, and SE into some of the Sunda Is and Taiwan. In the S the species ranged along the Nile Valley to Khartoum, and N of the Sahara in Africa, more orless following the continental coasts of S, E, and SE Asia. Within this range it was absent only from extremely dry deserts, e.g. the driest regions of Mongolia and in China W of Sichuan; and alpine zones, such as the high altitudes of Pamir and Tien Shan. In recent centuries, the range of S. scrofa has changed dramatically because of hunting and changes in available habitat. The species disappeared from the British Is in the 17" century, from Denmark in the 19" century, and was greatly reduced in range and numbers in the 20" century from areas as distant as Tunisia, Sudan, Germany, and Russia. Following these severe declines, there were some slight population recoveries in Russia, Italy, Spain, and Germany in the mid-20™ century, and natural and assisted range expansions in Denmark and Sweden. The species has also been inadvertently reintroduced in various locations in the Great Britain via escapees of mixed origin from commercial farming enterprises. Ex-S. scrofa stocks also occur as introduced feral populations in various other parts of the world, including Australia, New Zealand, the eastern Malay Archipelago, and in North, Central, and South America. In all of these areas they are now generally recognized as a major pest. in Suidae
Subspecies and Distribution. S. s. scrofa Linnaeus, 1758 — W Europe, from Denmark, Germany, Poland, and Czech Republic to N Italy and N Iberian Peninsula; possibly also Albania. The taxonomic status of animals in Austria, Switzerland, Slovenia, and Slovakia is unclear but presumably these populations are included in scrofa, as are the populations of Sweden, Finland, and the Baltic states. However, restocking of once depleted populations, for example in Italy, has likely involved the introduction and mixing of this subspecies with other subspecies, such as attila. S. s. affinis Gray, 1847 — S India and Sri Lanka. S. s. algirus Loche, 1867 — Tunisia, Algeria, and Morocco, on the coastal side of the mountains or in the low montane areas. S. s. attila Thomas, 1912 — Hungary, Ukraine, C & S Belarus, Romania, Moldova, and S Russia towards the N flank of the Caucasus, but not including the Transcaucasian countries of Georgia, Armenia, and Azerbaijan. The range possibly extends as far S as the Mesopotamian Delta in Iraq, in which case it would likely include W & SW Iran, and possibly E Turkey and Syria, where it borders with lybicus. Such a range could not be easily reconciled with a statement by Groves that "the difference between pigs from N and S of the Caucasus is quite striking; Transcaucasian boars are certainly not attila." This subspecies may also extend into C Asia and include Kazakhstan, Uzbekistan, and Turkmenistan, but no data exist to support this. S. s. baeticus Thomas, 1912 — originally described from Coto Donana, S Spain, and later merged with meridionalis; also S Portugal. Unless evidence is found that these Italian and Iberian populations are the relics of a much larger formerly contiguous range, this subspecies should be kept as distinct. S. s. coreanus Heude, 1897 — Korean Peninsula. S. s. eristatus Wagner, 1839 — Himalayas S to C India and E to Indochina (N of the Kra Isthmus). S. s. davidi Groves, 1981 — the arid zone from E Iran to Gujarat, including Pakistan and NW India, and perhaps N to Tajikistan. S. s. leucomystax Temminck, 1842 — main Is ofJapan (Honshu, Shikoku, Kyushu, Nakadori, Hiburijima, Tojima, Kushima, and other smaller Is). S. s. lybicus Gray, 1868 — Bulgaria, Greece, Turkey, Syria, Jordan, Israel, Palestine, in the past also in Lybia, and Egypt. The former Yugoslavia was included in its range, which would suggest that now Slovenia, Serbia, Croatia, Bosnia and Herzegovina, Montenegro, and Kosovo are within the range of this subspecies, although the exact boundaries are unclear. Pigs from Albania have been assigned to S. s. scrofa. S. s. majori De Beaux & Festa, 1927 — C & S Italian Peninsula. S. s. menidionalis Forsyth Major, 1882 — Corsica and Sardinia, with the proviso that the two populations are very likely to be introduced or feral. S. s. moupinensis Milne-Edwards, 1871 — China, S to Vietnam and W to Sichuan. S. s. nigripes Blanford, 1875 — the flanks of the Tianshan mountains in Kyrgyzstan and NW China (Xinjiang). An animal photographed in NE Iran (Golestan) looked like this subspecies. S. s. nukiuanus Kuroda, 1924 — Iriomote, Ishigaki, Okinawa, Tokunoshima, Amamioshima, and Kakerome Is in the Ryukyu chain in extreme S Japan, though some of these populations have hybridized with introduced domesticates. S. s. sibiricus Staffe, 1922 — Mongolia and Transbaikal (S & E of Lake Baikal). S. s. tawvanus Swinhoe, 1863 — Taiwan. S. s. ussuricus Heude, 1888 — far E Russia and the Manchurian region (China). Korean populations were previously included in this subspecies, but based on new evidence, the Korean taxon seems more similar to moupinensis. S. s. vittatus Boie, 1828 — Malay Peninsula, S of the Isthmus of Kra, the offshore islands of Terutai and Langkawi, Sumatra, Riau Archipelago, Java, Bali, and a range of smaller islands around these, including Babi, Bakong, Batam, Bawean, Bengkalis, Bintan, Bulan, Bunguran, Cuyo, Deli, Durian, Enggano, Galang, Jambongan, Karimon (Riau Is), Kundur, Lagong, Laut, Lingga, Lingung, Mapor, Moro Kecil, North Pagai, Nias, Panaitan, Payong, Penang, Pinie, Rupat, Siantan, Siberut, Simeulue, Singkep, Sugi, Sugi Bawa, Telibon, Tinggi, Tuangku, and the Tambelan Is. This species was originally present from the British Is in the extreme W, through Eurasia from S Scandinavia to S Siberia, extending as far E as Korea and Japan, and SE into some of the Sunda Is and Taiwan. In the S the species ranged along the Nile Valley to Khartoum, and N of the Sahara in Africa, more orless following the continental coasts of S, E, and SE Asia. Within this range it was absent only from extremely dry deserts, e.g. the driest regions of Mongolia and in China W of Sichuan; and alpine zones, such as the high altitudes of Pamir and Tien Shan. In recent centuries, the range of S. scrofa has changed dramatically because of hunting and changes in available habitat. The species disappeared from the British Is in the 17" century, from Denmark in the 19" century, and was greatly reduced in range and numbers in the 20" century from areas as distant as Tunisia, Sudan, Germany, and Russia. Following these severe declines, there were some slight population recoveries in Russia, Italy, Spain, and Germany in the mid-20™ century, and natural and assisted range expansions in Denmark and Sweden. The species has also been inadvertently reintroduced in various locations in the Great Britain via escapees of mixed origin from commercial farming enterprises. Ex-S. scrofa stocks also occur as introduced feral populations in various other parts of the world, including Australia, New Zealand, the eastern Malay Archipelago, and in North, Central, and South America. In all of these areas they are now generally recognized as a major pest.
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